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SMITHSONIAN INSTITUTION 
UNITED STATES NATIONAL MUSEUM 



PROCEEDINGS 



UNITED STATES NATIONAL MUSEUM 



VOLUME 103 

NUMBERS 3311-3337 




UNITED STATES 

GOVERNMENT PRINTING OFFICE 

WASHINGTON : 19S6 



ADVERTISEMENT 

The scientific publications of the National Museum include two 
series, known, respectively, as Proceedings and Bulletin. 

The Proceedings, begun in 1878, are intended primarily as a medium 
for the publication of original papers, based on the collections of the 
National Museum, that set forth newly acquired facts in biology, 
anthropology, and geology, with descriptions of new forms and 
revisions of limited groups. Copies of each paper, in pamphlet 
form, are distributed as published to libraries and scientific organiza- 
tions and to specialists and others interested in the different subjects. 

The dates at which these separate papers are published are recorded 
in the tables of contents of each of the volumes. 

The present volume is the hundred and third of this series. 

The Bulletin, the first of which was issued in 1875, consists of a 
series of separate publications comprising monographs of large 
zoological groups and other general systematic treatises (occasionally 
in several volumes), faunal works, reports of expeditions, catalogs of 
type specimens, special collections, and other material of similar 
nature. The majority of the volumes are octavo in size, but a quarto 
size has been adopted in a few instances in which large plates were 
regarded as indispensable. In the Bulletin series appear volumes 
under the heading Contributions from the United States National 
Herbarium, in octavo form, published by the National Museum since 
1902, which contain papers relating to the botanical collections of the 
Museum. 

Remington Kellogg, 
Director, United States National Museum. 



CONTENTS 

Pages 

Abbott, K. Tucker. Review of the Atlantic periwinkles 
Nodilittorina, Echininus, and Tedarius. Figures 55-57. 

Published March 23, 1954 449-464 

Arnett, Ross H., Jr. Beetles of the oedemerid genus 
Vasaces Champion. Figure 13. Published April 30, 

1953 87-94 

New species: Vasaces linearis, V. knulli, V. maculatus, V. elon- 
gaius, 

. A review of the beetle family Cephaloidae. 

Figure 20 and plate 5. Published May 15, 1953 .... 155-161 

. Supplement and corrections to J. A. Hyslop's 

Genotypes of the elaterid beetles of the world. Published 

April 14, 1955 599-619 

Banner, Albert H. A supplement to W. M. Tattersall's 
Review of the Mysidacea of the United States National 

Museum. Published July 8, 1954 575-583 

Berger, Andrew J. On the anatomy and relationships 
of glossy cuckoos of the genera Chrysococcyx, Lampro- 
morpha, and Chalcites. Figures 69-71. Published January 

19, 1955 585-597 

Blake, Doris Holmes. The chrysomelid beetles of the 
genus Strabala chevrolat. Figure 17. Published June 5, 

1953 121-134 

New species: Strabala rotunda, S. acuminata, S. colombiana, S. 

trinitatis. 
New subspecies: Strabala ambulans jamaicensis, S. a. puerto- 
ricensis, S. rufa floridana, S. acuminata teapensis, S. a. costa- 
ricensis. 

Carriker, M. a., Jr. Studies in Neotropical Mallophaga, 
XI: Bird lice of the suborder Amblycera, genus Denny us 
Neumann. Figures 63, 64. Published May 21, 1954 . 533-549 

New species: Dennyus brevicapitis, D. intonsus, D. rotundocapitis, 
D. brunneitorques, D. similis, D. spininotus, D. limbus. 

Cart WRIGHT, O. L. Scarabaeid beetles of the genus 
Bradycinettdus and closely related genera in the United 
States. Figures 14-16 and plates 3, 4. Published 
June 5, 1953 95-120 

New genera: Bolborhombus, Bolbocerastes. 

New species: Bradycinetulus rex, Bolbocerastes regalis, B. imperi- 

alis, Bolborhombus parvulus. 
New subspecies: Bolbocerastes imperialis kansanus. 

in 



IV PROCEEDmGS OF THE NATIONAL MUSEUM 

Carvalho, Jos6 C. M. Neotropical Miridae, LXIV: 
New bugs of the subfamily Cylapinae (Hemiptera). 
Figures 72-76 and plate 15. Published March 2, 1955 . 621-632 

New genera: Brachyfulvhis, Peritropoides. 

New species: Brachyfulvius chapmi, Peritroipoides annulatus, 

P. quadrinotatus, Vannius oculatus, Fulvms albonotatus, F. 

castaneoiis, F. ornatifrons. 

GuRNEY, Ashley B. Notes on the biology and immature 
stages of a cricket parasite of the genus Rhopalosoma. 
Figures 8, 9 and plate 1. Published March 10, 1953 . . 19-34 

. Distribution, general bionomics, and recogni- 
tion characters of two cocla'oaches recently established 
in the United States. Figure 10 and plate 2. Pub- 
lished March 10, 1953 39-56 

Hershkovitz, Philip. Mammals of northern Colombia, 
preliminary report No. 7: Tapirs (genus Tapirus), with 
a systematic review of American species. Figures 58- 
62. Published May 18, 1954 465-496 

Hyman, Libbie H. North American triclad Turbellaria, 
XIII: Three new cave planarians. Figures 65-68. 
Published June 14, 1954 563-573 

New species: Phagocata caver nicola, Sphalloplana georgiana, 
Speophila hoffmasteri. 

Kenk, Koman. The fresh-water triclads (Turbellaria) of 
Alaska. Figures 21-25 and plates 6-8. Published 
June 5, 1953 163-186 

New species: Phagocata nivea, Polycelis borealis, Dendrocoelopsis 
piriformis, D. alaskensis. 

Lachner, Ernest A. A revision of the goatfish genus Upe- 
neus with descriptions of two new species. Plates 13, 14. 

Published May 18, 1954 497-532 

New species: Upeneus asymmetricus, U. oligospilus. 

La Rivers, Ira. Two new naucorid bugs of the genus 

Ambrysus. Figure 1. Published February 12, 1953 . . 1-7 

New species: Ambrysus thermarum, A. bispimis. 

Lawrence, R. F. Two new scale-mite parasites of lizards. 

Figures 2-7. Published March 10, 1953 9-18 

New species: Geckobia keegani, G. philippinensis. 

McDermott, Frank A. Photuris hethaniensis, a new 1am- 

pyrid firefly. Published February 26, 1953 35-37 

New species: Photuris bethaniensis. 



CONTENTS V 

Morrison, J. P. E. The relationships of Old and New ^"^^ 

World melanians. Plate 11. Published April 20, 1954 . 357-394 

Pettibone, Marian H. Marine polychaete worms from 
Point Barrow, Alaska, with additional records from the 
North Atlantic and North Pacific. Figures 26-39. Pub- 
lished May 21, 1954 203-356 

ScHULTZ, Leonard P. Review of the Indo-Pacific anemone 
fishes, genus Amphiprion, with descriptions of two new 
species. Plates 9, 10. Published July 3, 1953 187-201 

New species: Amphiprion tricinctus Schultz and Welander, Am- 
phiprion mauritensis Schultz. 

AND Marshall, N. B. A review of the labrid fish 

genus Wetmorella with descriptions of new forms from the 
tropical Indo-Pacific. Figures 52-54 and plate 12. Pub- 
lished April 21, 1954 439-447 

New species: Wetmorella ocellata, W. albofasciata, W. triocellata. 
New subspecies; Wetmorella philippina bifasciata. 

Sturtevant, a. H. Nearctic flies of the family Periscelidae 
(Diptera) and certain Anthomyzidae referred to the family. 
Published June 16, 1954 551-561 

New species: Periscelis occidentalis, Cyamops imitata. 

White, Theodore E. Preliminary analysis of the fossil 
vertebrates of the Canyon Ferry Reservoir area. Figures 
40-51. Published May 28, 1954 395-438 

New genus: Kentrogomphios. 

New species: Kentrogomphios strophensis, Paradjidaumo spo- 
kanensis, Eumys cricetodontoides, E. latidens, E. spokanensis. 

Wirth, Willis W. Biting midges of the heleid genus Stilo- 
hezzia in North America. Figures 11, 12. Published 
May 15, 1953 57-85 

New species: Stilobezzia (Neosiilobezzia) stonei, S. (S.) beckae, S. 
(S). punctipes, S. (S.) sybleae, S. (S.) thomsenae. 

. American biting midges of the heleid genus Mono- 



helea. Figures 18, 19. Published June 17, 1953 .... 135-154 

New species: Monohelea (Monohelea) lanei, M. (M.) macfiei, M. 
(M.) texana, M. (M.) ornata, M. (M.) stonei, M. (M.) guianae, 
M. (M.) johannseni. 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 




Issued BMrlrvA- ScjSB ^y '^* 



SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 



Vol.103 Washington: 1953 No. 331 1 

TWO NEW NAUCORID BUGS OF THE GENUS AMBRYSUS 

By Ira La Rivers^ 

The following new Ambrysi were discovered after the manuscripts 
dealing with United States and Mexican species had been completed, 
and this paper is offered as a supplement to these treatments. Keys 
are included to allow the incorporation of the new species into the 
overall keys for the two regions. 

Family Naucoridae 

Genus Ambrysus Stdl, 1862 

Ambrysus thertnarum, new species 

Figure 1, &, c 

Description. — A small, pale species, nearly approaching Airibrysus 
fwnebris La Rivers, 1949, as the smallest species in the genus, and 
greatly resembling the latter in general conspectus; size 7.0-8.0 mm. 
long and 4.5-5.0 mm. wide. Dorsum bicolored, the scutellum and an- 
terior portion yellowish, the posterior area (hemelytra) brownish. 
Venter brownish, contrasting strongly with the yellow-white ap- 
pendages. 

Head glistening, yellow, slightly roughened and weakly punctate, 
protuberant before eyes and with the faintest suggestion of trunca- 
tion in front. Eyes blackish, slightly raised above general head sur- 

* Department of Biology, University of Nevada, Reno, Nev. 

228422—52 1 



2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

face when viewed posteriorly. Head ratios are : Total length to width 
(including eyes) 19 : 28 (68 percent) ; anterior distance between eyes to 
posterior distance 12: 17 (71 percent) ; anterior distance between eyes 
to inner eye-margin 12 : 11 ; posterior distance between eyes to greatest 
length of head posterior to this line 17 : 6 ( 35 percent) . 

Pronotum glistening, densely, shallowly punctate; color yellow to 
amber ; lateral edges smoothly rounded, nonserrate, occasionally with 
some weak pilosity. Curvature 14 percent (average 25:4), postero- 
lateral angles well rounded. Dorsal ratios are: Width between an- 
terior angles to greatest pronotal width 29: 51 (57 percent) ; median 
length to greatest width 17: 51 (33 percent) ; width between anterior 
angles to distance between anterior angle and posterior baseline of 
pronotum 29 : 25 (86 percent). 

Scutellum yellow-amber, shiny; shagreened with dense, shallow, 
white-rimmed punctation ; ratio of three sides 30 : 24 : 24. 

Hemelytra uniformly brownish, yellowing medially and in em- 
bolar region; shagreened as is scutellum. Embolium about normal 
for the genus, length-to- width ratio 29:9 (31 percent), rarely with 
some sparse, occasional marginal pilosity. Hemelytra moderately 
exposing lateral connexival edges, which latter have some prominent, 
short yellow pilosity. Hemelytra just attaining abdominal apex. 

Venter reddish brown, contrasting with the light-colored legs. 
Connexival posterolateral angles weakly acute-angulate, gradually 
enlarging posteriorly, not in the least spinose; connexival edges 
smooth, nonserrate, moderately pilose. Female subgenital plate 
apex simply concave, the concavity somewhat angular at its deepest 
point. Male genital process sharply right-angulate, tapering to apex, 
appearing much like a very thin foot, curving abruptly outward. 

Leg I yellow-white, darkening to light amber on tibia-tarsus; 
ratio of length to greatest width of femur 27: 16 (59 percent) ; com- 
bined tibia-tarsus, when closed, just attaining, or slightly surpassing, 
adjacent (proximal) end of femur. 

Leg II yellow ; ratio of femoral length to median width of ventral 
surface 25:5 (20 percent), length 1.9 mm.; tibia with distal end 
ventrally with one complete transverse terminal row of reddish spines 
set solidly across tip (remnants of the second, subterminal row, when 
complete and unbroken by use, extend about two-thirds of the width of 
tibia), ratio of length to width §2: 2 (10 percent), length 1.75 mm. 

Leg III yellow; ratio of length to femoral width 30: 5.5 (18 per- 
cent), length 2.1 mm. ; distal end of tibia ventrally with the same type 
of transverse spination as in mesotibia ; ratio of length to width 37 : 2 
(5 percent), length 3.0 mm. 

Tibia has the characteristic spinate appearance in contrast to the 
smoother micro-armed femora, with the long, silky, swimming hairs 
on metatibia contrasting with the less hirsute appearance of the 



TWO NEW NAUCORID BUGS — LA RIVERS 



mesotibia. All tarsi, except protarsi, 3 -segmented ; the first seg- 
ment minute ; the last two long and slender, terminating in two mod- 
erately curved claws. 

Type and paratopes.— VSKM. 60987. 

Type locality. — New Mexico, Sierra County, Hot Springs. 

Material examined. — Holotype (male), allotype, and four para- 
types, collected by Loew, September 11, 1874 (P. R, Uhler collection), 
in U. S. National Museum. 

Remarks. — This unique little species, discovered during the course 
of checking the Ambrysi in the National collection, is probably an 
offshoot, as far as present material can indicate, of Aimbrysus morTnon 
stock. The female subgenital plate is mormonoid in structure, and 
the variable 7normo7i-ty'pe male process could easily have given rise 





Figure 1. — Diagnostic characteristics of Ambrysus bispinus and A. thermarum: a. Apex 
of subgenital plate of A. bispinus, female holotype (visible only in ventral view), X22; 
b, apex of subgenital plate of A. thermarum, female allotype, X22; c, genital process on 
caudal edge of tergite V of A. thermarum, male holotype, X22; d, bispinate condition 
of left connexivum of abdominal sternite V of A. bispinus, female holotype, showing (1) 
anterior posterolateral angle and (2) posterior laterocaudal angle (ventral view, with 
caudal tip of abdomen oriented at the top of the field), X22. 

to the distinctive male process of A. thermarum, which has evolved 
toward the A. woodhuryi type to such an extent that, with only the 
male to examine, it would have been considered an aberrant A. wood- 
huryi. At present, it is the most definitely adapted thermal ecad of 
the A. mormon type, but as nothing is known of the structure of the 
type locality, it is impossible to say whether this seems to be due to 
complete isolation or not. 



4 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loe 

The following modification of my key to United States Ambrysi 
(La Rivers, 1951) will satisfactorily compare A. thermanmi with its 
closest relatives : 

8 (7). Female subgenital plate simply, broadly, and smoothly concave apically, 
the lateral bordering angles sharp; concavity, measured in terms of 
depth of concavity against length between bordering lateral angles, never 
less than 20 percent ; in clean material embolium nearly unicolorous, only 
very faintly lighter exteriorly, no contrasting colors present. 

buenoi Usinger, 1946 

Female subgenital plate, if simply concave, has (a) rounded lateral angles, 

or (b) if lateral angles are sharp, then depth of concavity is considerably 

less than 20 percent; in clean material embolium always sharply and 

contrastingly transversely bicolored, anterior two-thirds light yellow, 

posterior one-third blackish-brown 8A 

8A (8). Male genital process narrowing to apex, strongly, abruptly and almost 
right-angularly turned outward, giving the appearance of a long, 
attenuated foot with a rather sharp heel ; female subgenital plate 
simply concave at apex, somewhat angularly so. 

thermarum, new species 

Male genital process not definitely narrowing to apex, and not foot- or 

boot-shaped, although curving variably outward ; female subgenital 

plate simply and roundly concave at apex__ mormon Montandon, 1909 

Ambrysus bispinus, new species 

Figure 1, a, d 

Description. — A large, darkly mottled species, well rounded and con- 
vex; size 12.0-14.0 mm. long and 7.0-9.0 mm. wide. Dorsum not 
distinctly bicolored (i. e., anterior portion lighter than posterior area) , 
more or less uniformly mottled. Venter yellow or brownish, uni- 
colorous and lighter than dorsum. 

Head glistening, yellow-brown, smooth, weakly and vaguely punc- 
tate, slightly and broadly protuberant, and truncate before the eyes. 
Eyes blackish, nearly flush with plane surface of head when viewed 
posteriorly. Head ratios are : Total length to width (including eyes) 
34:54 (63 percent) ; anterior distance between eyes to posterior dis- 
tance 26 : 33 (79 percent) ; anterior distance between eyes to inner eye- 
margin 26 : 25 ; posterior distance between eyes to greatest length of 
head posterior to this line, 33 : 8 (22 percent) . 

Pronotum glistening, yellowish with brown mottling, sparsely 
punctulate, rugulose behind head; lateral edges smooth, nonserrate, 
rounded, on one specimen with traces of faint pilosity which appears 
to have been rubbed off. Curvature 10 percent (average 52:5); 
posterolateral angles well rounded. Dorsal ratios are: Width be- 
tween anterior angles to greatest pronotal width 53 : 73 (73 percent) ; 
median length to greatest width 39 : 73 (53 percent) ; width between 
anterior angles to distance between anterior angle and posterior base- 
line of pronotum 63 : 52. 



TWO NEW NAUCORID BUGS — LA RIVERS 5 

Scutellum glistening, yellow medially and on posterior angle, 
yellow along lateral edges, reddish brown elsewhere; shagreened with 
dense, coarse, white-spotted punctures ; ratio of three sides 70 : 50 : 50. 

Hemelytra mottled brown and yellow, glistening ; shagreened as is 
scutellum. Embolium slightly narrow for the genus, length-to-width 
ratio 60: 18 (30 percent), with no detectable marginal pilosity in the 
specimens examined. Hemelytra moderately exposing connexival 
margins, which latter have sparse, short pilosity, concentrated some- 
what at the posterolateral angles. Hemelytra just, to not quite, at- 
taining abdominal tip. 

Venter yellow to brown, the legs usually not markedly contrasting 
with the body in color. Connexival posterolateral angles acute-ang- 
ulate, short, gradually increasing in size posteriorly, not in the least 
spinose ; margins with some weak dentation, detectable only with con- 
siderable magnification, on all segments except I and II. Female 
segment V (the insect being oriented so that the abdominal tip is at 
the top of the field) possesses two posterolateral connexival processes ; 
the exterior posterolateral angle and an interior, prominent, blacker, 
rather blunt elongation of the caudal margin of segment V lying 
adjacent to the posterolateral angle. I have referred to this in other 
species as the laterocaudal coimexival angle. Tip of female sub- 
genital plate strongly and distinctively quadrisinuate, the lateral ter- 
minal angles long and rather sharp, but not extending as far caudad 
as the median, paired sinuosities which lie close together as a doubly 
rounded, blunt process longer than the lateral spines. Character of 
the male genital process unknown. 

Leg I amber ; ratio of length to greatest femoral width 48 : 31 
(65 percent) : combined tibia-tarsus, when closed, just attaining adja- 
cent (proximal) end of femur. 

Leg II amber; ratio of length to median width of femur 46:10 
(22 percent), length 3.4 mm.; tibia with distal end ventrally with 
one-and-one-half transverse rows of reddish spines, the terminal row 
complete, the subterminal row, as indicated above, only half complete : 
ratio of length to width 40 :4.5 (11 percent), length 3 mm. 

Leg III amber; ratio of length to median femoral width 57: 11 (19 
percent) , length 4.2 mm. ; tibia with same type of transverse terminal 
spination as in mesotibia, ratio of length to width 70: 5 (7 percent), 
length 5.2 mm. 

Tibia possesses the typical spinate appearance in contrast to the 
smooth femora ; long, golden, silky, swimming hairs are present on the 
metatibia and lacking on the mesotibia. All tarsi, except protarsi, are 
3-segmented, the first segment minute, the last two long and slender, 
terminating in two moderately curved claws. 

Type and paratype. — USNM 60988. 



6 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Type locality. — ^Mexico, Veracruz, Jalapa. 

Material examined. — ^Holotype (female) and a single female para- 
type from Mexico, Oaxaca, Camotlan, collected by W. S. Miller, April 
14, 1944, in the U. S. National Museum. 

Remarks. — In external appearance, A. bispinus is superficially in- 
distinguishable from A. cosmius and, as such, has probably shared 
the latter's confusion as A. signoreti Stal, 1862, in collections. How- 
ever, the structure of the female subgenital-plate apices is quite dif- 
ferent in the two species. In this respect, A. hispinus most closely 
resembles A. guttatipennis Stal, 1876, but while the subgenital plates 
might not be able, alone, to separate all variables of both species, the 
possession of primary and secondary connexival angles on abdominal 
segment V, with present material, sharply sets A. hispinus off from 
all other Ambrysi of the signoreti group. These two angles are more 
specifically referred to as the posterolateral and laterocaudal con- 
nexival angles, and the laterocaudal is lacking in all other known 
members of the group. It has shown up in other, distant sections 
of the genus, being known in A. parviceps Montandon, 1897, A. hun- 
gerfordi Usinger, 1946 (both in the pudicus group), and A. puncti- 
collis Stal, 1876 (a monotypic group). It is this character upon 
which the specific name of A. hispinus is based. In addition, the 
male genital process, when known, will furnish more evidence of the 
species' affiliations, either supplementing or diverting present indica- 
tions. 

The following couplets will enable the new species to be incorpo- 
rated in my key to Mexican Ambrysi now in press : 

24 (15). Female subgenital plate, viewed ventrally, asymmetrical on left side, 

where a prominent flap occupies the border beyond (laterad of) the 
left lateral angle ; right border scarcely produced. 

dilatus Montandon, 1910 

Female subgenital plate, viewed ventrally, not markedly or noticeably 

asymmetrical from one lateral border to the other 25 

25 (24). Female possessing both primary and secondary posterolateral connexi- 

val spines on abdominal sternite V (i. e., the primary angle being the 
posterolatei-al angle proper, the secondary angle being the laterocaudal 

angle) bispinus, new species 

Female lacking the secondary (laterocaudal) angle. 

remaining Mexican species of the signoreti group 



TWO NEW NAUCORID BUGS — LA RIVERS 7 

References 

La RrvEBS, Ira 

1949. A new species of Ambrysus from Death Valley, with notes on the 

genus in the United States (Hemiptera: Naucoridae). Bull. 
Southern California Acad. Sci., vol. 47 (1948), No. 3, pp. 103-110. 

1950. The meeting point of Ambrysus and Pelocoris in Nevada. Pan-Pacific 

Ent. vol. 26, No. 1, pp. 19-21. 

1951. A revision of the genus Ambrysus in the United States (Hemiptera : 

Naucoridae). Univ. California Publ. Ent., vol. 8, No. 7, pp. 277- 
338, illustr. 
— The Ambrysus of Mexico (Hemiptera: Naucoridae). Univ. Kansas 
Sci. Bull. (In press.) 

MONTANDON, A. L. 

1897. Hemiptera cryptocerata. — Sous-fam. Cryptocrieinae. Verh. zool.-bot. 

Ges. Wien, vol. 47, pp. 6-23. 
1909. Naucoridae. Descriptions d'especes nouvelles. Bull. Soc. Roum. Sci., 

Bucharest, vol. 18, No. 1, pp. 43-61. 
StAl, Carl. 

1876. Enumeratio Hemipterorum. Svenska Vetensk. Akad., vol. 14, pt. 5, 

No. 4. 
UsiNGER, Robert L. 

1946. Notes and descriptions of Ambrysus St&l with an account of the life 

history of Ambrysus mormon Montd. (Hemiptera: Naucoridae). 

Bull. Univ. Kansas, Sci. Bull., vol. 31, No. 1, pp. 185-210. 



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Vol. 103 Washington: 1953 No. 3312 



TWO NEW SCALE-MITE PARASITES OF LIZARDS 



By R. F. Lawrence ' 



Dr. G. W. Wharton, of Duke University, Durham, N. C, has kindly 
submitted to me for identification a number of scale mites from lizards 
of the families Gekkonidae and Iguanidae. The material, from the 
Philippine Islands, consists of one series taken from a lizard at Fort 
McKinley, Rizal, and another from the gecko Hemidactylus freTiatus 
in a house at Manila. Both series consist of two quite different species 
of Geckobia. These are described in this paper as new and both of 
them have been taken at the two localities mentioned above. There 
is thus good reason for assuming that the hosts of the two series are 
the same and that the "lizard" of Fort McKinley is identical with 
the gecko H. frenatus from which scale mites were taken at Manila. 
All these parasites were collected by Dr. H. L. Keegan, Third General 
Medical Laboratory, at Fort McKinley, Rizal, Philippine Islands. 

A further series, consisting of a large number of adults and larvae, 
were collected from the iguanid lizard Sceloporus undulatus^ at Dur- 
ham, N. C, by Dr. A. S. Pearse; these have been identified as Banks' 
New World pterygosomid Geckohiella texana. 

Several species of the scale mite parasite Geckobia may live on the 
same host; Hirst (1925, p. 173) has already pointed this out, saying, 
"When more than one form of Geckobia is found on the same host, 
one lives beneath the ventral scales and is flattened, being considerably 
wider than long and having scales instead of hairs on the venter. 
The second form occurring on the same host is usually to be discovered 
between the claw and pad of the toes, between the laminae of the pad, 
or between the toes themselves; this form is practically spherical in 
shape and has hairs instead of scales on the venter." Although there 

* Natal Museum, Pietermaritzburg, Union of South Africa. 

228383—53 9 



10 



PROCEEDINGS OF THE NATIONAL MUSEUM 



is no definitive information that the two strikingly different species 
of Geckobia here described actually illustrate Hirst's observation, it 
will probably be found that the two forms live on different parts of 
the body of the same host. 

Family Pterygosomidae 
Genus Geckobia Megnin, 1878 

Geckobia keegani, new species 

Figures 2, 3 

Material examined. — Three females, cotypes, from a lizard, Fort 
McKinley, Kizal, Philippine Islands, collected by H. L. Keegan, Octo- 
ber 1948, USNM 1931 ; also nine females bearing same locality data 




Figure 2. — Geckobia keegani, new species, dorsal surface. 



SCALE-MITE PARASITES — LAWRENCE 



11 




Figure 3. — Geckohia keegani, new species: a, Dorsal scute; h, peripheral dorsal hair; c, 
anterior ventral hair; d, posterior ventral hair. 

as the cotypes, and five females from Hemidactylus frenatus^ from 
Manila, Philippine Islands. 

Body almost circular; dorsal surface as in figure 2, with fairly 
long, almost parallel-sided hairs differing very little in length, but 
posteriormost slightly longer than anteriormost. Dorsal scute well 
defined, with about 20 hairs differing very little in general appear- 
ance and size from the remaining dorsal hairs, though a little shorter 
than the longest of these. 

Ventral surface with a density of hairs similar to that of the dorsal 
surface but reaching further back, almost to the anal field; the dis- 



12 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

crepancy in the length of these hairs far greater than in those of the 
dorsal surface, anteriormost a little less than half as long as posterior- 
most, latter a little shorter than the longest dorsal hairs. 

Mouth parts normal, pedipalps short and stout, both second and third 
segments above with a setif orm hair, that of the second weakly barbed 
and only a little stouter than that of the third, which is smooth. 

Legs equal-sized, rather small in proportion to the body. Pos- 
terior common coxa differing from most species of Gechobia in having 
two spurlike hairs on its anterior and only two, one, or even none 
on its posterior margin ; usually only three or four hairs, instead of 
the normal five, are found on this coxa and these are comparatively 
small and weak. The basal segments of the legs without spurlike 
hairs ventrally; but the second segment in all legs with a rather 
long, slender, and weakly barbed hair above. 

Dijnensions. — Width of body 0.603 mm., length (including mouth 
parts), 0.632 mm. 

Remarks. — The species closely resembles Gechohia simplex Hirst, 
1926, described from Hemidactylus leschenaulti., Madras, India, in 
the shape of the hairs of the dorsal surface and in the form of the 
dorsal scute. It differs from this species in having only about half 
the number of hairs on the dorsal scute, these being also more similar 
to the remaining body hairs in length and thickness than is the case 
in G. simplex. It further differs in there being very little difference 
in the shape of the dorsal hairs of the second and third palpal 
segments. 

Geckobia philippinensis, new species 

Figures 4, 5 

Material examined. — Two females, cotypes, from a lizard. Fort 
McKinley, Rizal, Philippine Islands, collected by H. L. Keegan, 
October 1948, USNM 1932. 

Body much wider than long, dorsal surface as in figure 4, with 9 or 
10 large, wide hairs in the area usually occupied by the dorsal scute, 
those in the middle of the body much smaller, while towards the pe- 
riphery posteriorly and laterally, the hairs become progressively more 
elongate and pointed. A single pair of centrally situated hairs much 
longer than the predominantly short hairs by which they are sur- 
rounded. Dorsal scute not defined. 

Ventral surface as in figure 5, a, which represents a section in the 
middle line of the body extending from its anterior to its posterior 
margin; the transition from the small type of cylindrical hair to 
scales of lanceolate form, sudden and without intermediate types of 
hair; the scales toward the posterior periphery becoming progres- 
sively more elongate and narrow. 



SCALE-MITE PARASITES — LAWRENCE 



13 




o 



14 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Mouth parts normal, pedipalps short, with rounded segments, sec- 
ond segment above with a stout plumose hair several times thicker 
than that of the third segment, which is smooth and setiform. 

Leg IV distinctly longer and a little stouter than the remaining 
ones; posterior common coxa as in figure 5, e, with five very large 
and distinct conical spurlike hairs; all legs with a fairly stout 





d 




Figure 5. — Geckobia philippinensis, new species: a. Hairs of ventral surface in midline of 
body; b, peripheral dorsal hair; c, small hair from middle of dorsum,'^(/, a scalelike ven- 
tral hair; e, posterior common coxa. 

plumose hair on the ventral surface of segments 2 and 3 and a 
long setose hair on the dorsal surface of segment 3, that of leg 
IV weakly plumose. 

Additional material examined. — Two females from the same host 
and locality as the types. Dr. Wliarton sent with his material the 
lizard host from which the Manila specimens were taken. From the 
body scales of this gecko, Hemidactylus fre^iatus^ I obtained a single 
female of the species described above. 



SCALE -MITE PARASITES — LAWRENCE 15 

Dimensions. — ^Width of body 0.400 mm., length (including mouth 
parts), 0.287 mm. 

According to Hirst's key (1925, p. 174) , this species would be brack- 
eted with Geckohia australis Hirst, 1926, but in width of body and 
general form of dorsal hairs and ventral scales, it seems to resemble 
more closely G. hindustanica Hirst, 1926, described from Hemidacty- 
lus leschenaulti., Madras, India. It differs from G. hindustanica in 
the complete absence of a dorsal scute and in the relatively larger size 
of leg IV, as well as in other details. 

Genus Geckobiella Hirst, 1917 

This genus was erected by Hirst (1917, p. 138) for the scale mites 
of Iguanid lizards, and thus far its members have been found to 
occur only on species of the genus Sceloporus. It would be of great 
interest to know what types of scale mites occur on the other numerous 
genera of the family Iguanidae in the New World. 

The eyes, which are not mentioned in the description of the genus, 
consist of a single pair situated in much the same position as those 
of Geckobia and Zomorobia., Lawrence, 1935, being located anteriorly 
near the lateral margin of the body. They are small, only a little 
larger han the ringlike sockets of the dorsal hairs (fig. 6, &). 

Geckobiella texana (Banks), 1905 

Figures 6, 7 

OecJcobia texana Banks, Proc. Ent. Soe. Washington, vol, 8, p. 134, 1905. 
OeckoMella texana (Banks), Hirst, Ann. Mag. Nat. Hist,, ser. 8, vol. 19, p. 
138, 1917, 

Material examined. — Numerous adults and larvae of this species 
collected by A. S. Pearse from Sceloporus undulatus at Durham, 
N. C, in January 1950. 

Adult female. — ^Hirst (1925, p. 200, fig. 19) figured only the ventral 
surface of this species, and the opportunity is now taken to give an 
illustration of the dorsal surface (fig. 6) . The dorsal hairs are much 
more numerous in some specimens than in others, there being probably 
a certain amount of variation in this respect. 

The mouth parts are longer and more robust than in most genera 
of Pterygosomidae ; the free portion of the peritremes reach to the 
distal end of the second palpal segment; the second palpal segment 
with a fairly long slender hair of equal thickness throughout, not 
pointed at the apex, with fine barbs ; dorsal hair of the third palpal 
segment longer, pointed apically and smooth or almost so ; fourth pal- 
pal segment with a single seta above, claw stout, conical, and somewhat 
curved. 



16 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figure 6. — Geckobiella texana (Banks): a, Dorsal surface of female; b, eye. 

Leg I only a little longer than IV, distinctly stouter than the re- 
maining legs ; segments 2 and 3 of all legs with a setif orm hair above, 
finely barbed. 

Adult male.—Th^ single specimen which appears to be an adult 
male is considerably smaller than the female and with far fewer hairs 



SCALE -MITE PARASITES — LAWRENCE 



17 





Figure 7. — Geckobiella texana (Banks): a. Dorsal surface of larva; h, penis (?). 

on the dorsal surface; otherwise it does not differ from the female; 
the chitinous genital structures are rather ill-defined, the large penis 
(?) (fig. 7,5), projecting slightly from the genital aperture at the 
posterior extremity of the body. 

Larva. — Dorsal surface with very few symmetrically disposed hairs, 
as in figure 7, a. Eyes present in the same position as the adult. Ven- 
tral surface entirely without hairs or setae except for a single seta at 
the base of legs I and III. Mouth parts closely resembling those of 
the adult, but the peritremes much shorter, reaching to a little below 



18 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

the base of the second palpal segment, and the claw of the palp more 
slender. Legs shorter than in the adult. 

Dimensions. — Female : Length of body 0.913 mm. ; width, 0.487 mm. 
Male : Length of body 0.652 mm. ; width, 0.326 mm. Larva : Length 
of body 0.740 mm. ; width, 0.413 mm. 

References 

BttEST, ARTHtm Stanley 

1917. On some new mites of the suborder Prostigmata living on lizards. 

Ann. Mag. Nat. Hist., ser, 8, vol. 19, p. 136. 
1925. On parasitic mites of the suborder Prostigmata (Trombidioidea), on 
lizards. Journ. Linnaean Soe. London, vol. 36, p. 173. 
Laweence, R. F. 

1935. The prostigmatic mites of South African lizards. Parasitology, vol. 27, 
No. 1, p. 6. 



U. S. SOVERNMENT PRINTING OFFICE: I9B3 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 




SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 



Vol. 103 Washington : 1953 No. 3313 

NOTES ON THE BIOLOGY AND IMMATURE STAGES OF A 
CRICKET PARASITE OF THE GENUS RHOPALOSOMA 



By Ashley B. Gurney ^ 



Introduction 



For more than three-quarters of a century wasps of the genus 
Rhopalosoma Cresson, 1865, have attracted the interest of hyme- 
nopterists because characters of the adult have left doubt regarding 
family relationships. Although the family Rhopalosomatidae ^ was 
proposed for the genus in 1896, there remains uncertainty as to what 
other genera are to be included in the family.^ Neither is it clear 
which of the related families is most closely allied to the Rhopalosoma- 
tidae. Biological studies of Rhopalosoma together with a critical 
examination of the immature stages may be helpful in clarifying these 
matters. Until now the only original published information concern- 
ing the biology and immature stages is that by Hood (1914) . He col- 
lected a larva which was reared to maturity and identified as R. poeyi 
Cresson, which is currently referred to R. nearcticum Brues. Hood's 
account is brief and, while it is a good general indication of the 
biology, contains no detailed morphological descriptions. The recent 
capture of a larva of Rhopalosoma at Falls Church, Va., has given 

^ Bureau of Entomology and Plant Quarantine, Agricultural Research Administration, 
U. S. Department of Agriculture. 

=" Originally given as Rhopalosomidae by Ashmead (18&6). An adult specimen of the 
genus has been illustrated by Westwood (1874, pi. 24, fig. 9). The species shown there is 
Rhopalosoma aenigmaticum (Westwood) of Hispaniola. 

8 Readers are referred to Brues and Melander (1932, p. 499), Brues (1943), and Krom- 
bein (1951). 



228353—53 



19 



20 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

impetus to the gathering of biological data, and the material at my 
disposal has also permitted morphological studies to be made on 
several of the immature stages. 

I am much indebted to Dr. Henry K. Townes, of the North Carolma 
State College (in absentia), for lending specimens of Rhojyalosoma 
and for supplying collecting records and giving helpful suggestions. 
My colleagues Dr. B. D. Burks, K. V. Krombein, and Dr. W. H. 
Anderson have also advised me generously. David G. Shappirio, of 
Harvard University, and Dr. T. H. Hubbell, of the University of 
Michigan, have kindly supplied data on parasitized crickets preserved 
at the University of Michigan Museum of Zoology. 

Biological observations 

The larva found at Falls Church ^ was in the fourth instar, attached 
to the left side of an adult female of Hapithus agitator Uhler. While 
1 was beating deciduous underbrush for psocids, on September 30, 
1950, the cricket was knocked into a collecting umbrella. The left 
hind leg was held at an unnatural angle with the body and the cricket 
was rather inactive and easily picked up. Leaves and other organic 
material were added to the jar in which the insect was placed, but the 
cricket was not seen to feed. The next day it scarcely changed its 
position, though the antennae occasionally moved. Meanwhile, the 
ventral surface of the parasite remained securely attached to the 
cricket and no movement was noticed. A 7 p.m. on the evening of 
October 2 the larva was beginning to shed its skin, as shown by the 
appearance of the white integument of the fifth instar where the 
fourth instar skin had split lengthwise along the middle of the 
dorsum. The cricket had then lost its foothold and was dying on its 
side, though its antennae still moved. 

By 8 : 30 p.m. the larva (which was entirely white except for traces 
of pale brown about the mandible) had left the dead cricket and had 
moved an inch or so away. The larva was placed in a pill box of soil 
where it gradually darkened to a creamy yellow color. On the day 
following the molt the larva made frequent twisting contractions as 
if trying to burrow. The head was directed downward most of the 
time and one silk strand coming from the mouth was noticed. The 
larva did not succeed in becoming completely covered with soil ; dur- 
ing the next day it was relatively inactive, with the head uppermost 



* Generic Identification of the parasite is primarily based on the very close agreement 
with the situation reported by Hood (1914). It may eventually be shown that more than 
one nominal form of Bhopalosoma inhabits the United States and that some specimens 
from the vicinity of Washington, D. C., are distinct from R. nearcticum, the type locality 
of which is Kissimee, Fla. 



CRICKET PARASITE — GURNEY 21 

and showing occasional movements of the mouth parts. By the morn- 
ing of October 5, it was decidedly quiescent and evidently dying, and 
by noon it had died and so was preserved in alcohol (pi. 1). If 
several inches of fine sand had been provided instead of heavy gar- 
den loam, perhaps the insect w^ould have entered the soil and suc- 
cessfully spun a cocoon. 

The present larva was attached to the cricket on the membrane 
between the terga and sterna of abdominal segments 1 to &. After 
the fifth instar had left, the cricket (pi. 1) bore three exuviae 
visible from above, representing the second, third, and fourth instars. 
In each the head was directed posteriorly and located farther posteri- 
orly on the cricket than that of the previous instar. The strongest 
single point of larval attachment was apparently just anterior to the 
parasite's modified anus ; there the attachment was quite secure. The 
larval body was noticeably curled, but the entire ventral surface was 
closely appressed to the cricket. The exuviae of larval instar 1 was 
situated transversely on the ventral surface of the cricket, in a groove 
just behind the left hind coxa. The head end of the skin was directed 
mesally on the cricket and extended almost to the sternal plate of the 
first abdominal segment, while the posterior end was beneath the ends 
of succeeding exuviae. Remnants of the eggshell occurred with the 
exuviae of the first instar. On the body of the cricket, in part be- 
neath the cast skins and also directly adjacent to them and extending 
briefly onto the left tegmen (front wing), was an accumulation of 
dried mucus or similar material. This was scalelike in appearance 
and I first mistook it for exuviae. The larval head was not inserted 
through the cricket's body wall, nor was there any definite opening, 
but the membrane where the last feeding mandibles (fourth instar) 
were applied showed signs of having been pierced. The points of 
contact for the mandibles of the second and third instars were covered 
with the exuviae of later instars. 

A second larva of RJiopalosom-a in the U. S. National Museum was 
collected on a last instar male nymph of Hapithus sp. at Marietta, Ga., 
August 9, 1947. The larva is in the fifth instar, though it probably 
was captured in an earlier stage since the host cricket is also preserved. 
The exuviae which were attached to the left side of the cricket agree 
with those of the Falls Church specimen so far as has been noted. 

The third larA^a of Ehopalosoina examined (fig. 9, e, f) was preserved 
in the fourth instar after having been collected on an adult female of 
Hapithus agitator at Fuguay, N. C, September 12, 1950, by Townes, 
Rabb, and Howden (North Carolina State College collection) (pi. 1). 
The head of the final feeding stage of the larva was on the cricket's 
fifth abdominal segment, with the mandibles applied to the mem- 
brane close to the spiracle and nearer the tergum than to the 



22 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

sternum. A crust of a dried secretion was present, as in the case of 
the Falls Church larva. Though a crust may have been associated 
with the Marietta, Ga., material, none was seen. The eggshell and 
first larval exuviae of each of the three larvae occurred immediately 
behind the hind coxa of the cricket. Likewise, the hind leg, of each 
cricket, adjacent to the parasite, was held outward as shown in the 
photographs. 

A fourth larva examined is one of several which Dr. Townes has 
told me of finding on Hapithus and Orocharis. Of this particular 
larva he has written, "On September 12, 1950, 1 collected (at Fuguay, 
N. C.) a large Orocharis nymph with a one-third grown larva on it. 
I kept the cricket in a jar and its parasite prospered. On September 
17 I found the remains of the demolished nymph in the jar and the 
Rhopalosoma larva, now creamy white with a large head and pro- 
jecting mandibles, busily hitching itself along with its mandibles. 
It was very restless, so I put it in some damp sand. The larva 
promptly worked into the sand and came to rest in one corner of the 
jar, under the sand. Three days later it had completed an elongate, 
castaneous cocoon." 

The cocoon was kept indoors by Dr. Townes, at a temperature in 
the neighborhood of 55° F. In late February he mailed it to me and 
suggested that dissection to determine whether pupation had occurred 
might be advisable. The parchmentlike cocoon was 14 millimeters 
long, about 5 millimeters in diameter, and rounded at the ends. The 
inner surface was a lighter chestnut color than the outside and scarcely 
any individual threads of silk were apparent except on the outside. 
When the cocoon was opened on February 23, the larva was doubled 
up in the middle of it with about one-fourth of the cocoon's length 
empty at each end. Pupation had not occurred, nor had there been 
any casting of larval skins or passing of fecal material, but the larva 
appeared perfectly healthy. After preservation it was 6 millimeters 
long. The mandibles bore no teeth. 

The original observations of Hood (1914) were based on a larva 
taken at Plummer's Island, Md., October 6, 1912. The host, Orocharis 
saltator Uliler, was quite active when found, as it was said to be 
"scampering over the forest floor as rapidly as a large abdominal 
protuberance and a nearly functionless hind leg would permit." The 
anterior end of the parasite was at the host's eighth abdominal seg- 
ment. Tliis specimen grew rapidly, left the cricket three days after 
capture and entered the soil, emerging indoors as an adult about 
March 1, 1913. The information Hood gave concerning the exuviae 
and the quiescent state of the host just prior to the parasite's final 
visible molt agrees fully with my findings. The cocoon from which the 
adult emerged has recently been opened and the fifth instar exuviae 



CRICKET PARASITE — GURNEY 23 

were relaxed and studied. The mandibles of the exuviae differ some- 
what in shape from those of the Falls Church larva and they bear no 
teeth. 

In the University of Michigan Museum of Zoology are at least 20 
pinned cricket specimens bearing parasitic larvae presumed to be of 
one or more species of Rhopalosoina. I have not examined them, 
but detailed host data have been furnished me, together with the state- 
ment that the larvae vary considerably in size and that all are brown. 
The specimens were collected by Dr. T. H. Hubbell and several asso- 
ciated collectors and former students. The hosts, with the number 
of parasitized specimens. States where collected, and generalized 
dates of collection are as follows : Hapithus agitator agitator Uhler, 
seven specimens, southeastern Indiana, central Missouri, Mississippi, 
Tennessee, all collected in the last half of August ; H. agitator quad- 
ratus Scudder, four specimens, Florida, late July to early September; 
H. hrevipennis Saussure, four specimens, Florida, late July to mid- 
November ; Orocharis saUator Uhler, four specimens, July 4 to July 
29. The host associated with an additional larva from Florida is 
not recorded. These records represent the parasitism of this type 
occurring among several hundreds of Hapithus and Orocharis 
specimens. 

The specimen of Hapithus agitator on which the Falls Church larva 
was parasitic was preserved in alcohol and later dissected to determine 
the condition of its internal organs. Scarcely any traces of food 
were in the digestive tract, none of recent ingestion. Keproductive 
organs were evidently intact, but there were no well-developed eggs 
or other signs of normal functioning of the organs. 

The impression created was that the cricket had been weakened 
to a point where the drain on its vitality finally caused death, though 
there had been no mechanical destruction of organs. 

Various details of the biology remain to be clarified, but the main 
outline seems evident. Upon hatching, the larva probably crawls part 
way from the eggshell and starts its development in a more or less 
transverse position. At the first molt the larva apparently shifts 
the anterior part of the body so that it extends longitudinally along 
the side of the cricket, though keeping the same place of anal attach- 
ment. The third and fourth instars each move the head posteriorly 
on the cricket at molting time. Sharp mandibles occur in the second, 
third, and fourth instars, and these apparently pierce the cricket's 
integument and permit fluids to be taken into the mouth. Definite 
mouth parts of the first instar have not been seen, but may occur. 
The fifth instar does not feed but is an inactive stage that leaves the 
host and spins a cocoon in the soil. Pupation is evidently delayed, 
perhaps until near emergence time of the adult. 



24 PROCEEDINGS OF THE NATIONAL MUSEUM vol.103 

Regarding habits of Rhopalosoma adults, Dr. Townes has furnished 
the following notes : "I have seen the adults on several occasions and 
usually was fortunate to catch them when seen. I should say that 
in the Upper and Lower Austral Zones of the Atlantic States they 
are widespread and not uncommon in the right habitat. This appears 
(.0 be places of dense, shrubby vegetation where there is considerable 
humidity, as along stream bottoms and seashores, or, in other words, 
the same sorts of places where Hapithus and Orochuris abound. The 
adults may be flushed from the bushes. They fly up and alight again 
like an ophionine ichneumonid. They may be distinguished from 
ophionines in flight only by a slightly stockier appearance. At Long 
Beach, North Carolina, at twilight on July 9, 1949, I saw about 10 
all together in normal flight, cruising in a rather slow but erratic 
manner just over the beach shrubbery, again like an ophionine ich- 
neumonid in its twilight flight. They were flying until no longer 
visible in the gathering dark. I could catch only two of these and 
they were both males." 

The following collection dates accompanying specimens identified 
as Rhopalosoma nearcticum are available as an indication of when 
adults are active : Maryland : Mayo Beach, September 14, 1947, female 
(Townes); Berlin, July 19, 1932, female. North Carolina: Long 
Beach, July 9, 1949, two males (Townes) ; Wallace, August 2, 1949, 
female (Townes). South Carolina: Greenville, September 2, 1940, 
male (H. and M. Townes). Georgia: Stone Mountain, August 13, 
1949, female (Fattig) ; Atlanta, August 11, 1946, female. Florida : 
About a dozen specimens, essentially all reported by Brues (1943), 
ranging from June 9 to August 13, but mostly collected in June and 
July. 

In addition to the foregoing States, nearcticum has been recorded 
from Missouri and Kentucky by Krombein (1951). 

The known hosts of Rhopalosoma in the United States are Orocharis 
saltator Uhler and Hapithus agitator Uhler. Judging from the rec- 
ords of parasitized crickets preserved in the University of Michigan 
Museum of Zoology, Hapithus h'revipennis Saussure appears to be a 
host also. The distribution of 0. saltator, as stated b}^ Hebard (1931, 
p. 217) , covers the entire southeastern United States, extending north- 
ward to southeastern Pennsylvania, Illinois, and Missouri, and un- 
doubtedly to eastern Texas in the Southwest. In southern Florida a 
primarily West Indian species, Orocharis gryUodes (Pallas) , occurs. 
H. agitator occurs as far northeast as Long Island, and otherwise is 
largely comparable to O. saltator in distribution (Blatchley, 1920, 
p. 740; Hebard, 1931, 1938; Rehn and Hebard, 1916, pp. 308-310). 
Two geographic subspecies are recognized, the southeastern one, H. 
agitator quadratus Scudder, living from the Atlantic Coast to Browns- 



CRICKET PARASITE — GURNEY 25 

ville, Tex., and in the East apparently merging with typical H. agita- 
tor in central Georgia and along tlie "fall line" in North Carolina. H. 
hrevipennis inhabits Florida, Georgia, and Louisiana, and doubtless 
other southeastern States. It may be that any of the crickets men- 
tioned above are satisfactory hosts of Rhopalosoma. In southeastern 
Pennsylvania adults of both Orocharis and Hapithus begin to appear 
by early August, and in all of the more northern States the crickets 
pass the winter as eggs, and adults persist until frost time in the fall. 
Blatchley (1920) reported finding hibernating adults of E. agitator 
in Florida and suggested that they represented a fall generation. The 
subfamily Eneopterinae, to which Eapithus and Orocharis belong, is 
primarily tropical, and various genera are well represented in the 
Neotropical Region, from which Brues (1943) has recognized four 
species of Rhopalosoma. 

Morphological descriptions 

Mature larva {fifth instar from Falls Churchy Va.). — General shape 
as in plate 1; strongly mandibulate; legs entirely absent; abdomi- 
nal segments moderately well demarked, less so posteriorly, those be- 
yond segment 8 not readily separated; tAVO thoracic and eight 
abdominal pairs of spiracles. 

Head in frontal view as in figure 8, / ; occipital border broadly and 
evenly rounded (asymmetrical in the figure because it is bordered by 
uneven cervical folds) ; vertex and frons not individually demarked, 
the integument shiny, slightly irregular, not noticeably punctate ; no 
frontal arms evident; paired conspicuous spots (ey), presumed to be 
vestigial eyes, just above base of clypeus ; antenna reduced to sensory 
pit with traces of facetlike division, situated well to the side of frons, 
no apparent projecting sensilla; a presumed sensory area represented 
by about nine poorly defined, pigmented spots (sa), with tiny seta at 
lateral margin of group; frons and vertex with very sparse, short 
setae, grouped as illustrated; six strong clypeal setae; labrum con- 
spicuously bilobed, each lobe with two major setae, lateral and an- 
terior margins with microsetae, an irregiilar row of pits across each 
lobe, some bearing short setae; mandibles heavily sclerotized, apex 
prominent and narrowly rounded, basal third swollen laterally and 
with a single seta, biting margin thin, demarked from main mandib- 
ular structure by groove, and sharply but delicately toothed. (Left 
mandible with four well-spaced teeth along middle third of biting 
margin, followed by closely set group of four; right mandible with 
two teeth, others perhaps lost by breakage.) Maxilla with several 
medium-sized setae as illustrated, apex near tip of mandible sharply 
rounded and with asperities, maxillary palpus barrel shaped, galea 



26 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

equally elongate but of smaller diameter ; labium with palpi subequal 
to galeae, area of sericteries with two angular projections posterior to 
weakly sclerotized nonsegmented elongate appendages of uncertain 
homology; apparent opening of silk duct (sd) between bases of latter 
appendages. 

Anterior spiracle on posterior third of prothorax; second spiracle 
barely anterior to groove separating meso- and metathorax; abdominal 
spiracles on segments 1 to 8, slightly more dorsad than thoracic ones, 
situated near anterior margin of the respective segments ; each spiracle 
unspecialized so far as visible under 72-power magnification, consist- 
ing of circular disk with central aperture (internal structures not 
studied) ; no setae on thorax or abdomen apparent; no apparent spe- 
cializations for crawling or clinging; vestigial anus transverse, at 
posterior margin of segment 10. 

Coloration : General color creamy yellow, darker on head (con- 
spicuous black areas on body shown in photograph are necrotic spots 
developed at time of death). Vertex, frons, and clypeus yellowish; 
eye spots, sensory areas, antennae, and pits at bases of setae pale brown- 
ish ; a darkened area anterior to sensory area ; a somewhat darkened 
indefinite band across base of clypeus connecting bases of mandibles ; 
labrum colorless, except for brown setae and pits ; mandibles marked 
as illustrated with rich, dark brown ; maxillary and labial palpi and 
galeae dark brown; labium white, elongate appendages in area of 
sericteries tinged with yellowish ; spiracles yellowish ; no special pig- 
mented areas about nonfunctional anus or elsewhere on thorax and 
abdomen. 

Measurements (in millimeters) : Length 8 (would actually be 
slightly longer if body not curved); greatest width of body 3.2; 
greatest width of head capsule 1.3. 

Comments: Fifth instar from Marietta, Ga., more robust than 
larva from Falls Church, Va. ; setae on frons and vertex smaller, 
scarcely noticeable; teeth on mandibles lacking. Measurements (in 
millimeters) : Body length 8.5 ; greatest width 4 ; greatest width of 
head capsule 1.6. 

Penultmiate stage larva {fourth instar from Fuguay^ N . C). — Gen- 
eral appearance typically hymenopterif orm (fig. 9, e, /) ; head capsule 
and mouth parts not so highly developed as in fifth instar ; legless ; 10 
pairs of spiracles ; modified anus apparently specialized as a hold-fast 
organ. 

Head capsule (fig. 8, c) rather masklike in contrast with that of 
fifth instar, integument very rough and irregularly rugose, a conspicu- 
ous tranverse ridge (epistoma) dorsad of labrum; two pairs of blunt 
protuberances on upper part of face, the median pair (mt) broad and 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 103 PLATE 1 




u = 



£1 



5=< 




V a; 

•-2 ~ 



5p 



DO 




< 



P 



CRICKET PARASITE — GURNET 



27 



labs 



mxs 




FiQUEE 8.— The genus Rhopalosoma Oresson, 1865: o, Mouthparts of fourth Instar exuviae; 6, external view 
of sixth abdominal spiracle of fourth instar exuviae; c, front view of head, fourth instar alcoholic specimen; 
d, antenna of fourth instar exuviae; e, anal region, fourth instar exuviae; /, front view of head, fifth instar 
alcoholic specimen; (c made from Fuguay, N. O., specimen, all others from Falls Church, Va., material). 



28 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 108 

irregularly rounded, lateral pair (It) narrow and evenly rounded; an- 
tennal socket (an, fig. 9, e) circular, minute, traces of sensilla scarcely 
visible under 72-power magnification; labrum membranous except 
mesally and at lateral extremities, strongly bilobed, each lobe bearing 
two small setae about equidistant from center and a minute seta near 
anteromesal margin ; exposed apical fourth of overlapping mandibles 
triangular, the apex sharply acute, about seven sharp teeth ; approxi- 
mate basal third of each mandible exposed near end of epistoma, bear- 
ing a single, minute seta ; a long tapering apodeme extending dorso- 
laterally within head capsule; maxillary sclerome (mxs) anterior to 
base of mandible, shaped as illustrated ; labial sclerome (labs) an ir- 
regular, incomplete circle. 

Abdomen with 10 apparent segments; spiracles located as in fifth 
instar ; entire dorsal area glabrous ; venter of prothorax and cervical 
region bearing minute asperities ; dorsal and lateral integument ap- 
pearing as if covered with closely appressed scales (see later descrip- 
tion of exuviae) ; eight abdominal segments with ventral, transverse 
groups of small platelets ; hold-fast organ at posterior margin of seg- 
ment 10, transverse, weakly crescent-shaped, with short membranous 
flap extending posteriorly. 

Coloration: General color dark blackish-brown; ventral region 
(unstippled areas on fig. 9, e, /) unpigmented; a white middorsal line 
(where split occurs at subsequent molt) extending entire length of 
body ; head considerably darker than thorax or abdomen ; region of 
frons lighter colored than main epicranial region, two darker pig- 
mented spots (ps) anterior to median tubercles, a wide longitudinal 
white stripe on frons, interrupted on anterior portion of vertex and 
resumed as narrow line on occiput; labrum unpigmented except at 
lateral extremities, on a slender transverse basal median sclerite, and 
mesally at the junction of the lobes; mandibles and scleromes of max- 
illae and labium pigmented with brown as illustrated. Abdomen 
with paired unpigmented spots dorsad of spiracles 3 to 8, and in a 
similar position on abdominal segment 9 (which bears no spiracle) ; 
transversely grouped platelets of ventral surface pale brown, also clos- 
ing sclerites of hold-fast organ and immediately adjacent membrane. 

Measurements (in millimeters) : Length of body 5.7; greatest 
width of body 2.2 ; greatest width of head capsule 1.4. 

Comments: Slide-mounted preparations of the cast skin of the 
fourth instar from Falls Church, Va., permitted structures to be 
observed in much greater magnification than was possible with the 
preserved alcoholic larva from Fuguay, N. C. While no significant 
differences in these individual fourth stage larvae apparently occur, 
it is important that the features illustrated and described be attributed 
to the correct specimens. 



CRICKET PARASITE GURNEY 



29 








d cc <:^o©'^^^20 



'^^::>z> c^ 





hg--^ 



Figure 9— The genus Rhopalosoma Cresson, 1865: a. Section from lateral area of thorax, showing closely 
grouped platelet structure, fourth instar exuviae; 6, integumental asperities on ventral surface of pro- 
thorax, fourth instar exuviae; c, d, specialized platelets on ventral surface of fourth and fifth abdominal 
segments, respectively, fourth instar exuviae; e, f, ventral and lateral views, respectively, of fourth instar 
alcoholic specimen; (e,f, from Fuguay, N. C, specimen, others from Falls Church, Va., material). 



an— antenna 
cv — closing valves 
ep — epistoma 
es— esophagus 
ey— eye 
ga— galea 
hg— hind gut 



labs— labial sclerome 
labr — labrum 
Ip— labial palpus 
It— lateral tubercle 
md — mandible 
mdc — mandibular condyle 
mt— median tubercle 



mxp— maxillary palpus 
mxs — maxillary scleroma 

ps— pigment spots 

sa — sensory area 

sd— opening of silk gland 
or sericteries 

tr— trachael ring 

vp— ventral platelets 



30 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

Exuviae split lengthwise on dorsum, from epistoma to tenth ab- 
dominal segment; presumed antenna consisting of circular socket 
(jBg. 8, d) with three tiny unpigmented circles in center, each bearing 
a central spot of pigment; mouth parts (fig. 8, a) with mandibles 
the most characteristic and best preserved of the parts ; mandible thin, 
much flattened, biting margin with large apical tooth and about 10 
acute smaller teeth in a single row along apical third, an emargination 
a little posterior to the teeth, portion of margin overlaid by labrum 
scarcely pigmented, a well-sclerotized and pigmented region at base 
of mesal margin; posterior margins of mandible with a prominent 
black ventral condyle, a small seta near base of long lateral apodeme 
(presumed mesal apodeme for muscle attachment missing in available 
material) ; labrum largely membranous, with irregular transverse 
wrinkles, a constricted pigmented sclerite mesally, about five subapical 
minute pits arranged transversely on each lateral half; a poorly 
defined, somewhat striated organ leading posteriorly to labial sclerome 
from base of labrum, dorsad of remnants of apparent esophagus 
(es) ; labial and maxillary scleromes showing somewhat different 
shape in flattened aspect than in natural position on alcoholic larva. 

Integument of most of thorax and abdomen smooth, microscopically 
marked into platelets of different shapes on various areas of the 
body, averaging transversely elongate ventrally, irregularly quadrate 
(fig. 9, a) dorsally on thorax and anterior half of abdomen; ventral 
region of neck and prothorax bearing minute asperities of varied 
character (fig. 9, J) , some setiform, others in form of sharp rugosities ; 
less developed asperities on ventral surface of abdomen; specialized 
platelets on venter of abdominal segments 1 to 8 (poorly developed 
on 1 to 3 and 8, well developed on 4 to 7), of irregular grouping on 4 
and 5 (fig. 9, c), others transverse (fig. 9, d). Spiracles each sur- 
rounded by narrow irregular area of membrane, the external portion 
a circular flattened disk with small central opening ; trachea tapering 
to spiracular opening, with internal ring (tr, fig. 8, &) which may 
function as closing apparatus. Modified anus (fig. 8, e) with well- 
sclerotized and pigmented closing valves (cv), integument at lateral 
extremities wrinkled, that posterior and anterior to valves moderately 
pigmented, a short section of membrane on internal surface an appar- 
ent remnant of hind gut (hg) . 

Exuviae of the first three instars are very closely appressed to each 
other along the posterior half of their abdominal sterna, so much so 
that they can scarcely be separated without injury to the specimens. 
The closest attachment appears to be a little anterior to the modified 
anal opening (hold-fast organ) . The platelets comprising the micro- 
structure of the inteis:ument on the lateral and dorsal surfaces of 



CRICKET PARASITE GURNET 31 

thorax and abdomen, which in the fourth instar vary from tightly 
grouped on the side of the thorax (fig. 9, a) to narrowly separated on 
the abdomen, are much less closely grouped in the third and second 
instars, while they are so distant and reduced in size in the first instar 
as to resemble very widely spaced spiculelike spots of pigment. 
Mandibles of the third and second instars roughly resemble those 
of the fourth, and bear sharp teeth, there being at least six to eight 
in the third and apparently fewer in the second. The epicranium of 
the first instar has about two pairs of rather large areas of darker 
pigment, but no mouth parts have been observed, though they may 
have escaped detection. There are 10 pairs of spiracles in the third 
and second instars, with the pre-spiracular tracheal ring poorly defined. 
Only six pairs of spiracles have been found on the exuviae of the first 
instar, and the posterior one is poorly developed. Each of the three 
early instars is brown, darker on the head. The venter is pale in 
the third and second instars. Approximate lengths of the exuviae of 
the Falls Chuch, Va., larva are as follows (in millimeters) : Fourth 
6.5 ; third 3.6 ; second 2.3 ; first 1.2. Those of the Marietta, Ga., larva 
are almost identical, but the Fuguay, N. C, larva is larger, the three 
exuviae measuring as follows : Third 4.4 ; second 2.6 ; first 1.3. 

Eggs apparently are broadly elongate-oval; fragments are about 
1.4 millimeters long. 

A basic paper on the larval head of parasitic Hymenoptera is that 
of Vance and Smith (1933). Among the more specialized papers, 
those of G. C. Wheeler (1943) and Beirne (1941) have aided me most 
while studying the anatomy of fifth- and fourth-instar heads, re- 
spectively, of Rhopalosoma. The book by Clausen (1940) is a val- 
uable aid toward understanding the biology and morphology of 
related families, and it should be consulted for references to the im- 
portant work of J. C. Bridwell, F. A. Fenton, R. C. L. Perkins, F. X. 
Williams, F. van Emden, and others. 

Summary and discussion 

Among the biological data additional to those brought out by Hood 
(1914) is the fact that Ha'pithus agitator is an acceptable host for 
Rhopalosoma in the eastern United States. The effect of the parasite 
on the internal organs of the host, unfortunately based on only one 
dissected cricket, has been described. Details concerning the position 
of the parasite egg and various larval stages have been given. Notes 
on adult habits, essentially lacking previously, make it clear that in 
parts of the Southeast specimens of Rhopalosoma are not nearly so 
scarce as has been supposed. Like many another rarity in collections, 
these insects can doubtless be taken in numbers by those familiar with 



32 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

their appearance, time of occurrence, and favored habitats. It is note- 
worthy that on a single day, September 12, 1950, Dr. Townes and his 
associates found two larvae, one on Hapithus^ the other on Orocharis. 
Specimens belonging to the University of Michigan show that on three 
other occasions, once in Indiana and twice in Florida, a collector has 
found two instances of parasitism in a single day. 

There remain many unanswered questions concerning the biology 
of the species of Rhopalosoma, some of which may require controlled 
cage studies. Stages earlier than the fourth instar are still known 
from exuviae and fragments of eggshell, and first instar specimens 
are especially desirable for morphological study. 

More study material, including specimens from northern and south- 
ern States, may show that more than one form of Rho'palosoma occurs 
in the United States. Adults of nearcticum were distinguished from 
their Neotropical congeners by Brues (1943) on rather minor morpho- 
logical features, and differences correlated with geographic distribu- 
tion might be better represented in larvae than adults. Of the four 
fifth-instar specimens examined, the ones from Georgia and North 
Carolina and the exuviae of Hood's Maryland specimen lack teeth on 
the mandibles, while the larva from Virginia apparently has a full 
complement of teeth on the left mandible but only a few basal ones on 
the right mandible. It is not known whether occasional breakage of 
teeth occurs while the larvae are burrowing in soil prior to cocoon 
formation. Some breakage of the teeth of the Virginia larva may 
have resulted from my manipulation of the mandibles with needles 
before I realized the existence of teeth. The three fifth-stage larvae 
have been measured for maximum head width and length of mandibles. 
The ratio of head width to length of mandible is 1.68, 1.86, and 2.0 for 
the larvae from Virginia, North Carolina, and Georgia, respectively. 

Brief examination of a few examples of other families has sug- 
gested certain characters which deserve careful attention in subse- 
quent work of a comparative nature. A chrysidid larva {Ghrysis) 
has the second thoracic spiracle located behind the groove separating 
meso- and metathorax, while the corresponding spiracle of Rhopalo- 
soma is slightly in front of the groove. Unlike the condition in 
Rhopalosoma^ the second spiracle of a tiphiid larva (Oosila) is smaller 
than adjacent ones. This tiphiid larva has two maxillary appendages, 
and there are two prominent angular projections on the labium in the 
area of the sericteries. As more details of rhopalosomatid life histories 
become available, comparisons with the situation in the Tiphiidae, 
such as has been discussed by Clausen, King, and Teranishi (1927), 
will perhaps be instructive. 

A larva of one of the Larridae {Larra analis Fabricius, which at- 
tacks mole crickets) has the head proportionately smaller than 



CRICKET PARASITE — GURNEY 33 

Rhopalosoma^ but as in the latter the maxilla and labium have two 
pairs of appendages each. A conspicuous difference is that the man- 
dible of Larva has a single preapical tooth in the incisor area, and 
basally on a lower lever there are two heavy molar teeth, or "pros- 
thecae." The mesal pair of elongate labial appendages of RhopalosoTna 
has apparent homalogues in this larrid. 

The mouth parts of those few larvae of Bethylidae and Dryinidae 
that I have seen are much reduced or otherwise specialized. Illustra- 
tions of dryinid larvae on the bodies of leafhoppers, such as those 
of Haupt (1916)^ show that the parasite larva sometimes remains 
associated with its exuviae in a manner suggestive of Rhopalosoma. 
The genus Harpagocryptus Perkins, based on a species that Perkins 
(1908) described from Queensland, is of interest in this connection 
because the larva was said to form "a sac on the sides of the abdomen 
of small crickets (Trigonidiidae)." Although originally referred by 
Perkins to the Dryinidae, Bischoff (1927, p. 407) called Harpago- 
cryptus a bethylid, and Reid (1939) suggested that it may be a psam- 
mocharid. As with Rhopalosoma^ a knowledge of the immature stages 
of Harpagocryptus might prove of great value when adequate com- 
parative studies eventually are possible. 

References 

AsHMEAD, William H. 

1896 Rhopalosomidae, a new family of fossorial wasps. Proc. Ent. Soc. 
Washington, vol. 3, pp. 303-309. 
Betkne, Bryan P. 

1941 A consideration of the cephalic structures and spiracles of the final 
instar larvae of the Ichneumonidae (Hym.). Trans. Soc. British 
Ent., vol. 7, pp. 123-190, 31 figs. 
Bischoff, H. 

1927 Biologic der Hymenopteren. 598 pp., 224 figs. Berlin. 
Blatchlet, W. S. 

1920 Orthoptera of Northeastern America. 784 pp., 246 figs. Indianapolis. 
Brites, Charles T. 

1921 Correlation of taxonomic affinities with food habits in Hymenoptera, 

with special reference to parasitism. Amer. Naturalist, vol. 55, pp. 
134-164. 
1943 The American species of Rhopalosoma. Ann. Ent. Soc. Amer., vol. 36, 
pp. 310-318, 9 figs. 
Brues, Charles T., and Melandeb, A. L. 

1932 Classification of insects. Bull, Mus. Comp. ZooL, vol. 73, pp. 1-672, 
1121 figs. 
Clausen, Curtis P. 

1940 Entomophagous insects. 688 pp., 257 figs. 
Clausen, Curtis P., King, J. L,, and Tebanishi, Cho 

1927 The parasites of PopilUa japonica in Japan and Chosen (Korea), 
and their introduction into the United States. U. S. Dep. Agr. Dep. 
Bull. 1429, 1-55 pp., 1 pi., 35 figs. 



34 PROCEEDmGS OF THE NATIONAL MXTSEUM vol. io3 

CUSHMAN, R. A. 

1926 Some types of parasitism among the Ichneumonidae. Proc. Ent. Soc. 
Wasliington, vol. 28, pp. 29-51, 51 figs. 
Haupt, H. 

1916 Beitrage zur Kenntnis der Cicadinenfeinde. Zeitschr. Wiss. Insekt- 
enb., vol. 12, pp. 217-223, fig. 13. 
Hebabd, Morgan 

1931 The Orthoptera of Kansas, Proc. Acad. Nat. Sci. Philadelphia, vol. 83, 
pp. 119-227, 2 figs. 

1938 Where and when to find the Orthoptera of Pennsylvania, with notes 

on the species which in distribution reach nearest this state. Ent. 
News, vol. 49, pp. 97-103. 
Hood, J. Douglas 

1914 Notes on the life history of Rhopalosoma poeyi Cresson. Proc. Ent. 
Soc. Washington, vol. 15 (1913), pp. 145-147, fig. 
Krombein, K. V. 

1951 [Rhopalosomatidae] in Muesebeck, et al., Hymenoptera of America 
north of Mexico. U. S. Dep. Agr. Monogr. 2, pp. 773-774. 
Perkins, R. C. L. 

1908 Some remarkable Australian Hymenoptera. Proc. Hawaiian Ent. 
Soc, vol. 2, pp. 27-35. 
IlEHN, J. A. G., AND Hebabd, Morgan 

1916 Studies in the Dermaptera and Orthoptera of the Coastal Plain and 
Piedmont region of the southeastern United States. Proc. Acad. 
Nat. Sci. Philadelphia, vol. 68, pp. 87-314, 3 pis. 
Reid, J, A. 

1939 On the relationship of the hymenopterous genus Olixon and its allies, 

to the Pompilidae (Hym.). Proc. Ent. Soc. London, ser. B, vol. 8, pp. 
95-102, 2 figs. 
Vance, A. M., and Smith, H. D. 

1933 The larval head of parasitic Hymenoptera and nomenclature of its 
parts. Ann. Ent. Soc. Amer., vol. 26, pp. 86-94, 8 figs. 
Westwood, J, O. 

1874 Thesaurus entomologicus Oxoniensis. 205 pp., 40 pis. 
Wheeler, George C. 

1943 The larvae of the army ants. Ann. Ent. Soc. Amer., vol. 36, pp. 

319-332, 2 pis., 2 figs. 
1948 The larvae of the fungus-growing ants. Amer. Midi. Nat, vol. 40, pp. 
664-689, 3 pis., 3 figs. 
Wheeler, William Morton 

1919 The parasitic Aculeata, a study in evolution. Proc. Amer. Philos. 
Soc, vol. 58, pp. 1-40. 



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Vol. 103 Washington: 1953 No. 3314 

PHOTURIS BETHANIENSIS, A NEW LAMPYRID FIREFLY 



By Frank A. McDermott 



In August 1949, at Bethany Beach in southern Delaware, some ob- 
servations were made on the few lampyrids then still flying. A single 
specimen of Photinus pyralis (Linnaeus, 1767) and one of Photuris 
versicolor (Fabricius, 1798) were observed, and a very few instances 
of a double greenish flash were noted over grass in a large vacant area 
at the north end of the town and perhaps 1,000 feet from the ocean 
front. The insects giving this flash were not captured, however, in 
spite of much pursuit, but another insect was seen to fly into the grass 
while giving the frequently repeated, rather dim flash characteristic of 
females of Photuris. This one was taken and proved to be a rather 
small female Photuris with a distribution of the black pigmentation on 
the pronotum different from any previously seen. Previous visits 
to Bethany Beach earlier in the year had yielded nothing notable, ex- 
cept the relative scarcity of lampyrids. In July 1951 I again spent a 
few days at Bethany Beach, and again saw the double greenish flash. 
This was considerably later than the usual period of prevalence of P. 
versicolor., and it seemed probable that this flash was that of a different 
species. Several males were taken, and one female, the latter unfor- 
tunately lost by accident, but it may be noted that the females are 
apparently relatively scarce. 

It differs from other photurids so far found in Delaware by the 
relatively small size, the distribution of the pronotal pigmentation, 
and the rather distinctly different flash of the male. The pronotal 
black pigmentation, while rather variable, has a distinct tendency to 
be enlarged toward the apical margin, and in none of the specimens 
did this spot show the forms characteristic of those of P. versicolor., 

228423—53 35 



36 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loa 

P. lucricrescens Barber, 1951, or P. hebes Barber, 1951, all of which, 
had been collected during the same summer in northern Delaware. 
The characteristic male flash is much like that of Photinus consangui- 
neus LeConte, 1851, two short, bright coruscations separated by a 
distinct interval perhaps somewhat longer than that of a single corus- 
cation, but is quite greenish as compared with the yellow light of 
consanguineus. Occasionally the second coruscation may be much less 
bright than the first, even with the same specimen, and a few appar- 
ently gave only a single flash but were indistinguishable from those 
giving the double flash. 

The first male specimens were taken on July 11, 1951, over the same 
field where the single female was taken in 1949, and where they were 
still scarce, but two days later a large colony was found flying over the 
vegetation, largely bayberry, on both sides of the road leading south 
from the town, at about two miles. Trips to areas a mile or so inland 
from the beach yielded only Photinus pyralis^ here very abundant at 
the time, and an occasional stray Photuris versicolor^ so the new species 
seems to be a rather strictly seacoast, almost a sand-dune, form. It 
has not been seen in several visits along the southern New Jersey coast, 
particularly at Stone Harbor, but it would be somewhat surprising if 
it were not present there. 

In view of these circumstances, this insect is here described from 
the males collected in 1951, and the single female of 1949, as a new 
species. 

Photuris bethaniensis, new species 

Description. — A rather small lampyrid for a Photuris, 9.0-10.75 
mm. long by 3.5-4.0 mm. wide, with a black pronotal mark tending 
to be densest tow^ard the apical edge. The characteristic male flash is 
two greenish coruscations separated by a short interA^al. 

Head: Width across eyes 2.1-2.45 mm. in male, 1.8 mm. in female; 
eye length 1.15-1.3 mm. in male, 0.85 mm. in female. Frons ivory to 
yellow, about 0.8-1.0 mm. wide; interocular margins usually rather 
divergent. Mandibles fairly large, 0.65-0.8 mm. across in closed posi- 
tion; proportionately larger in the female. Maxillary palpi dark 
brown, apical joint conoidal ; labial palpi lighter brown, apical joint 
mitten-shaped. Clypeus dark brown, more or less tridentate. 

Antennae: Dark brown, 5.0-6.25 mm. long in male, 4.25 mm. in 
female ; joints may all have pale bases, or this may be limited to basal 
three joints in the female. 

Pronottjm: Scutate, sometimes extended apically, 2.0-2.5 by 
2.55-3.0 mm. ; rounded posterior angles may or may not be appreciably 
produced ; red pigmentation may extend to the basal edge ; black spot 
of variable shape, and characteristically mainly toward the apex, or 



NEW LAMPYEID FIREFLY — McDERMOTT 37 

at least the larger portion of it apical, the longitudinal portion being 
usually merely an irregular and sometimes interrupted line, and but 
little developed along the basal margin. 

SCUTELLUM AND MESONOTAL PLATES I White Or yelloW. 

Elytra : 7.0-8.0 by 1.6-2.0 mm. ; ground color brown to nearly 
black; on each a white to yellow vitta, oblique, usually extending to 
about three-quarters the elytral length ; wide yellowish lateral borders 
which give a slightly oval contour; narrow sutural borders. 

Ventral thoeacic surface : Brown. 

Tekglfes : Tergites 1 to 6 brown ; tergite 7 lighter or only partially 
inf uscate ; tergite 8 pale to white. 

Sternites : 2 to 5 usually brown, sometimes becoming darker in this 
order; sternite 5 may be yellow; sternites 6 and 7 luminous in the 
male ; and sternite 8 with a median posterior point, which may be very 
short. The luminous area in the female differs from the pattern seen 
in versicolor^ being a transverse bilobed area medially very narrow, on 
segment 6, and a relatively large elliptical area on segment 7. 

Legs : Long ; tibial spurs long, pattern 0-2-2. 

Aedeagus : In the one dissected, 2.25 mm. long, identical in appear- 
ance with that of P. lucicrescens and the other photurids described by 
Barber.^ 

Type aifid paratypes. — USNM 61469. 

Type locality. — Bethany Beach, Del. 

Material examined. — Holotype (male) and five paratypes, collected 
by McDermott at type locality, July 11, 1951; paratype (female), col- 
lected by McDermott at type locality, August 1949. 

Remarks. — Photuris hethaniensis is probably a seacoast form; it 
has not been noted elsewhere. The flash suggests P. fairchildi Barber, 
1951, but the latter is a larger species with narrow elytral borders. 



* Barber, H. S. North American fireflies of the genus Photuris. Smithsonian Misc. Coll., 
vol. 117, No. 1, pp. 1-58, 3 figs., 1951. 



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Vol. 103 Washington: 1953 No. 3315 

DISTRIBUTION, GENERAL BIONOMICS, AND RECOGNITION 
CHARACTERS OF TWO COCKROACHES RECENTLY ESTAB- 
LISHED IN THE UNITED STATES 



By Ashley B. Gurney^ 



Introduction 



In contrast to the large numbers of cockroach species that occur 
in a great variety of natural habitats in most tropical countries, only 
about 55 are now recorded from the entire United States. With the 
exception of those occurring in a few localities, mainly in the South- 
western States, our roaches are well known to entomologists. In the 
northern third of the country there are few native roaches living in 
fields or woodlands, and most of the better-known species are estab- 
lished adventives which are rather closely associated with human 
habitations, having been brought there by artificial means. The recent 
establishment in the Northeastern States of two potentially important 
roaches new to this country is therefore of decided interest. 

Beginning in 1948 and continuing each subsequent year, specimens 
of Ectohius livens (Turton), the spotted Mediterranean roach,^ have 
been collected at Falmouth, Mass., under outdoor conditions, coming 
to houses at night and also occurring on fresh vegetables. Because 
of these habits and the wide natural occurrence of this roach in Europe 
and adjacent areas, it seems quite probable that a slow spread in the 
United States may take place, and that from time to time questions 
regarding its identity and importance will arise. 

1 Bureau of Entomology and Plant Quarantine, Agricultural Research Administration, 
U. S. Department of Agriculture. 

2 No generally accepted common name seems to be applied to E. livens in Europe, and 
this name is considered suitable for American use in view of the species' appearance and 
the region of its dominant Old World occurrence. 

39 

228354— 53— —1 



40 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

A second species not previously known to breed in the United States 
is LeiLCophaea maderae (Fabricius), the Madeira roach. Unlike the 
spotted Mediterranean roach, L. maderae is well known because it is 
frequently transported with articles of commerce in tropical countries. 
It has many times been intercepted by quarantine inspectors at 
United States ports and has occasionally been carried to inland cities, 
where it failed to survive. In the autum of 1950 it was found to 
be established in the basements of New York City buildings occupied 
by people who had come from Puerto Rico. Although its native 
home is evidently West Africa, it apparently was introduced to the 
West Indies more than 100 years ago. The Madeira roach could 
probably live outdoors in tropical Florida and perhaps at the other 
southern extremities of the United States, but it is more likely to be 
important as a domestic pest in the well-heated buildings of our larger 
cities. One has only to recall the increasing importance of the brown- 
banded roach Supella supellectiliuTii (Serville) in this country during 
comparatively recent years to realize the potential importance of 
Leucophaea maderae. Apparently a native of Africa that has been 
spread by commerce to most of the warmer parts of the world, S. 
supellectilium was first reported from the United States in 1903, based 
on specimens from Key West, Fla. It was not noted as a pest here 
until found in a private home in Nebraska in 1929, but since then 
has rapidly gained recognition as one of our principal pest roaches 
(see Back, 1937, and Gould and Deay, 1940) . 

This paper is intended to aid in the identification of these two 
roaches and to supply such available information on distribution and 
habits as is likely to be helpful to subsequent studies of these species 
in the United States. A third species, Nauphoeta cinerea (Olivier), 
was found to be established in Florida after this manuscript was 
prepared. It is briefly noted on page 46. 

Genus Ectohius Stephens, 1835 
Ectobius livens (Turton), 1800? (spotted Mediterranean roach) 

Figure 10, a-d: Plate 2, Figures 3-6 

Blatta livida Fabricius, Entomologia systematicae, vol. 2, p. 10, 1793. (Pre- 
occupied by Blatta livida De Geer, Memoirs pour servir a I'histoire des In- 
sectes, vol. 3, p. 538, pi. 44, fig. 6, 1773.) 

Blatta livens Turton. A general system of nature . . . , vol. 2, p. 529, 1800? 
(New name for Blatta livida Fabricius.) 

EctoMus lividus Fabricius, Stephens, Illustrations of British entomology, Mandi- 
bulata, vol. 6, p. 48, 1835. 

Ectohius livens (Turton) , Ramme, Mitt. Zool. Mus. Berlin, vol. 27, pp. 34-45, pi. 10, 
fig. 3, 1951. 

Nomenclatural coTnments. — ^Until 1951 Ectobius lividus was usually 
applied to this species. The name E. livens has been resurrected by 



COCKROACHES NEW TO UNITED STATES — GURNEY 41 

Kamme (1951, p. 34) who says that Dr. K. Princis of Lund, Sweden, 
informed him that livida Fabricius was preoccupied and that Turton 
proposed livens on page 526, vol. 2, of his 1806 edition of Linne's Sys- 
tema Naturae. Turton published two editions of an English transla- 
tion of the thirteenth (Gmelin) edition of this work. These editions 
are listed in the 1903-1915 Catalogue of the Books, Manuscripts, Maps 
and Drawings in the British Museum (Natural History) (vol. 3, 1910, 
p. 1128). I have verified their dates and contents by an examination 
of the entire 7-volume 1806 edition in the Library of Congress and of 
volume 2 of the 4-volume first edition in the library of the U. S. Na- 
tional Museum. The first Turton edition appeared from 1800 to 1802. 
Volume 2, containing the genus Blatta^ is dated 1800 and is reasonably 
sure to have appeared no later than 1802. The second Turton edition 
appeared in 1806, the first 4 volumes being re-issues of the former 
edition with slight changes of the title pages. On the page indicated 
by Ramme (p. 526) Forftcula livida^ an earwig, is treated. 

Prior to Ramme's monograph, the identities of the British species of 
Ectohius were confused. Thus the species treated by Lucas (1920, p. 
78) as pers-picillaris Herbst is in reality livens. This has been clarified 
by Blair (1934, 1935). Princis (1936) has explained that a Lapland 
record of lividus (i. e., livens) actually was based on material of 
lapponicus (Limiaeus). Hebard (1943, p. 12) states that Australian 
records of lividus are incorrect. Taxonomists now differentiate the 
species of Ectobius on rather minute characters, especially on the basis 
of the dorsal abdominal gland and genitalia of males. Ramme ( 1949 ) 
has described E. siculus from Sicily, differing from livens in having 
black legs and antennae. E. finoti Chopard of Algeria (see Chopard, 
1943, p. 21) is much like livens, having the small tegminal spots between 
the veins and with the dorsal gland bearing a rounded tubercle. The 
genus EctoMus (genotype: lappoQiicns) in a modern sense is much 
more restricted than formerly. In the older literature, species of 
EctoMus were reported from many parts of the world. Certain of 
these are now known to belong to other genera. For instance, many 
years ago several Nearctic roaches were referred to Ectobius. Rehn 
(1931, pp. 306-374) and Hebard (1943) have discussed the generic 
limits of Ectobius: Rehn (p. 308) has explained the nature of the 
toothed tarsal claws ; the teeth were evidently overlooked by Hebard 
(p. 12). 

Recognition features. — The most important features enabling rec- 
ognition of Ectobius livens are: (1) Small size, over-all length about 
8 to 9 millimeters ; (2) small, dark spots on the veins of the front wing 
(tegmen) ; (3) conspicuous intercalated triangle at apex of wing (itr, 
fig. 10, b) ', (4) claws of unequal length, the posterior claw of each 
pair much longer than anterior one; (5) in males the characteristic 
shape of the dorsal abdominal glandular depression and the sub- 



42 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

genital plate (fig. 10, «, c) ; (6) conspicuous spines on ventroposterior 
margin of each femur. 

Of the foregoing features, the second and fifth are specific for the 
separation of this roach from other species occurring in the United 
States and the others are highly confirmatory when in combination. 

The native Nearctic species most likely to be mistaken ^ for E. livens 
are Chorisoneura texensis Saussure and Zehntner, of Texas and the 
Southeast, and members of the genus Cariblatta. Both have claws of 
equal length, and Cariblatta lacks a well-developed, intercalated tri- 
angle. C. texensis has a large intercalated triangle, but has no con- 
spicuous spines along the posterior margins of the femora. The best- 
known outdoor roaches of the Eastern States belong to the genus Par- 
coblatta; all are larger than livens and lack an intercalated triangle. 

Identification of the Massachusetts specimens of livens was aided 
by comparisons made at the Academy of Natural Sciences of Phila- 
delphia with authentic French and Swiss material, given to the Acad- 
emy by the Swiss entomologist Henri de Saussure. 

Description. — Adults : General form as in plate 2, figure 6 ; tegmina 
and wings covering abdomen ; width between eyes on vertex about 1^ 
times length of first antennal segment (female), slightly less in male; 
all femora with conspicuous spines on ventroposterior margins ; ven- 
troposterior margin of front femur with three or four well-spaced, 
strong spines along part slightly basad of middle, the more apical one 
often smaller than others, followed by 10 to 15 small, delicate spines 
(some of which are frequently broken) extending in a regular row 
nearly to apex, a long curved apical spine closely preceded by a straight 
subapical one about three- fourths as long ; tarsal segments one to four 
each with a tiny, round pulvillus (membranous pad) ; segment 5 with 
large, broad apical arolium between conspicuously unequal tarsal 
claws, anterior claw about two-thirds length of posterior one, longer 
claw with about four spinelike inner teeth along basal half, teeth on 
smaller claw poorly developed. Tegmen with costal veins simple, 
cubital veins (discoidal sectors) strongly oblique, apex rather sharply 
rounded. Wing with distinctive intercalated triangle (itr, fig. 10, 

General color pale yellowish; tegmina and marginal area of pro- 
notum transparent, marked with reddish brown spots, on the tegmen 
these spots located on the veins ; eyes dark brown to black ; face yellow 
to reddish orange ; vertex and upper part of face usually with several 
spots of darker orange; coxae and femora unspotted; tibiae with 
dark brown at bases of some spines; venter of abdomen with poorly 

*The standard taxonomie reference on Nearctic Blattarla Is that of Hebard (1917). A 
recent key to genera is by John W. H. Rehn (1950). The European species of EctoMus 
were reviewed by Ramme (1923). 



COCKROACHES NEW TO UNITED STATES — GURNEY 43 

developed longitudinal rows of dark brown siibmarginal and median 
spots. 

Measurements (in millimeters) of representative specimens: Over- 
all length, including folded tegmina, female 9, male 8.5; pronotum, 
female 2.1, male 1.9 ; tegmen, female 7, male 7.2. Width of pronotum, 
female 3, male 2.9. No significant size variation has been noted. 

Nymphs : Nymphs (pi. 2, figs. 3, 4) ranging in body length from 
2.5 to 4.5 millimeters have been examined. They are yellow, with 
conspicuous dark reddish brown spots on the dorsal surface and on the 
vertex and upper part of the face. Dark submarginal longitudinal 
streaks appear on the thorax. The venter of the abdomen is dark 
brown submarginally and medially. 

Oothecae: Two oothecae have been examined (pi. 2, fig. 5). They 
are dark brown and measure 2.3 millimeters in length. The convex 
margin is weakly crenulate and there are faint transverse indications 
of the eggs contained. 

Distribution. — Three American localities are known for Ectobius 
livens, all in Massachusetts: Falmouth, Manomet, and Plymouth. 
Falmouth, the first site discovered here, is situated near Woods Hole 
and the Cape Cod Canal, on the southern side of the base of Cape Cod. 
In October 1951, after this manuscript had been prepared, I ex- 
amined 3 males and 4 females of livens collected at Manomet, Plym- 
outh County, Mass., by Mrs. S. P. Graeff and submitted by Dr. Ells- 
worth H. Wheeler of the University of Massachusetts. Manomet is 
on the coast of Cape Cod, about 20 miles north of Falmouth. The 
specimens were collected before August 13, 1951, apparently just 
prior to that date. Mrs. Graeff wrote, "Our summer cottage is pretty 
well overrun with them. Painters told me they were even on the roof. 
The cottage is in the woods and the underbrush is close. I think this 
is the third summer we have had them. I suspect they like damp 
spots, they scurry under shingles." 

An adult of livens was submitted July 28, 1952, by a Plymouth 
housewife who reported that her house and shrubbery were becoming 
infested. 

This roach is widely distributed in the southern and central portions 
of western Europe, occurring in Germany, the Netherlands, Belgium, 
France, Switzerland, Italy, Portugal, Spain, and southern England. 
It has also been reported as far eastward in the Mediterranean area 
as Asia Minor, but the records I have seen were published prior to 
the modern work dating from Ramme's 1923 monograph of Ectobius 
and are not dependable. Chopard (1943, p. 21) has reported it from 
Algeria and Tunisia. I have found no records of its occurrence in 
the Azores. Fabricius originally described the species from France, 
and, in the absence of type material, Eamme in 1923 designated a male 



44 PROCEEDINGS OF THE NATIONAL MUSEUIVI vol. io3 

neotype from Montpellier in southern France. Two varieties of 
livens have been recognized, chopardi Adelung described from France 
in 1916, and minor Kamme described from the Island of Elba in 1923. 
These varieties are based primarily on the amount of dark color on 
the lower surface of the abdomen, minor being quite dark except near 
the lateral margins, only the submarginal dark spots being well de- 
veloped in chopardi^ and the submarginal spots being poorly or moder- 
ately developed in typical livens. The species of Ectohius are well 
known for color variation, the color of tegmina and abdomen often 
being recessive (pale) or intensive (dark). In view of this situation, 
it remains to be proved whether chopardi and minor are valid sub- 
species or if they are forms without real significance. The center of 
distribution for the genus Ectohius^ as outlined by Rehn (1931, pp. 
314—318), was apparently the Mediterranean Basin, with a secondary 
center developed in Africa south of the Sahara Desert. 

Biology. — The original Massachusetts specimens of Ecto'hius livens 
examined consist of 2 males, 16 females, 9 nymphs, and 2 oothecae. 
The males were taken June 21, 1948, and the other specimens in late 
August and September 1950.^ It may be significant that males were 
found only in the spring and early summer. Lucas (1928) reported 
that in England males of E. lapponicus disappeared about midsum- 
mer, the females persisting longer than males, and nymphs appearing 
in late summer and fall preparatory to hibernation. No detailed 
life history studies of livens in Europe have come to my attention. 
Brown (1952) has reported observations on the related E. panzeri 
Stephens in England, which occurs there mainly on sand dunes near 
the coast. Adults die in the fall, the males disappearing first, but 
nymphs overwinter and do not reach full maturity until early August 
of the following year. 

The Falmouth collections were made on a farm located beside a 
small salt river about a mile from the shore. Mr. Flint found speci- 
mens in a large variety of situations, specimens occurring most con- 
sistently beneath loose lichens on oak trees, and crawling on trees and 
houses at night — the latter in the vicinity of lights. Some were 
found under baskets, in buckets, or on Swiss chard and other vege- 
tables, or were taken by general sweeping (see Flint, 1951). 

In England E. livens occurs only in the extreme southern counties — 
there found on trees, among bracken ferns, under dead leaves, and 
flying actively in hot sunshine (Lucas, 1920, p. 80; Burr, 1936, p. 44). 
Chopard ( 1947, p. 37) says livens is very common in the French woods, 
and he gives a colored illustration (pi. 2, fig. 28) . 

■* AU of this material was collected by Oliver S. Flint, Jr., a student at the University 
of Massachusetts. I am much indebted to Mr. Flint for the enthusiasm, with which he 
sought additional material once the character of his Initial captures was pointed out to 
him, as well as for his diligence in seeking clues to the origin of the introduction. 



COCKROACHES NEW TO UNITED STATES — GURNEY 45 

Source of introduction. — The Massachusetts colony of Ectohius 
livens is probably the result of an accidental introduction from western 
Europe or the Mediterranean area. A good many people of Portu- 
guese ancestry live in Falmouth and the vicinity, and visits to Portu- 
gal and the Azores are frequent. Seeds and occasional shipments 
of fruit are brought back, but quarantine regulations prevent wide- 
spread movements of plant products. Since livens occurs in Portugal, 
as reported by Seabra (1942, p. 19), the possibility of an introduc- 
tion from that part of the Mediterranean seems the most likely. 

Probable importance. — Ectohius livens is not likely to develop into 
a major pest or to live in buildings except occasionally. As a nuis- 
ance pest in the vicinity of gardens and dwellings, it may be expected 
to attract attention at intervals and to be submitted to entomologists 
for identification and advice, as is now true of male specimens of 
Parcohlatta which are attracted to lights and which occasionally re- 
main in houses for short periods. 

So far as known, species of Ectohius are not economically impor- 
tant in Europe. E. lapponicus has often been stated to be a pest of 
dried fish in Swedish Lappland, but Gaunitz (1935, 1936) found 
that laffonicus does not occur indoors and probably feeds on 
vegetable matter. 

Genus Leucophaea Brunner, 1865 
Leucophaea maderae (Fabricius), 1793 (Madeira roach) 

Figure 10, e-h; Plate 2, Figxjbes 1, 2 

Blatta maderae Fabricius, Entomologia systematicae, vol. 2, p. 6, 1793. 
Leucophaea maderae (Fabricius), Rehn, Trans. Amer. Ent. Soc, vol. 29, p. 283, 
1908. 

Nomenclatural comments. — The name Rhyparohia maderae is cur- 
rently used by certain workers, especially Europeans, as exemplified 
by Chopard (1943, p. 45) in his monograph of North African Orthop- 
tera. Leucophaea was proposed by Brunner (1865, pp. 272, 278) 
as a subgenus of Panchlora.^ with four included species. One of the 
four, maderae^ was designated type of Leucophaea by Rehn (1903, 
p. 282). Rhyparohia was proposed by Krauss (1892, p. 165), mono- 
typic for maderae. Thus Rhyparohia is an isogenotypic synonym of 
Leucophaea. The use of Rhyparohia was doubtless encouraged by the 
action of Kirby (1904, pp. 150-151) who, evidently unaware of Rehn's 
1903 designation, designated Blatta surinamensis Linnaeus type of 
Leucophaea. As explained by Hebard (1917, p. 309), Kirby also ap- 
parently objected to the name Pycnoscelus Scudder, 1862, to which 
surinamensis belongs, because it was originally based on an immature 
specimen. Kirby's latter view is not supported by the rules of nomen- 



46 PROCEEDINGS OF THE NATIONAL MUSEUM yol. los 

clature but, regardless of that aspect of the matter, the validity of 
Leucophaea^ with maderae its type, is clear. 

Recognition features. — The most important features in the recog- 
nition of Leucophaea maderae are : (1) Large size, overall length about 
38 to 53 millimeters; (2) pronotal shape, and general color markings 
as illustrated (pi. 2, figs. 1, 2) ; (3) no strong spines along posterior 
margins of femora except one short apical spine on each hind and 
middle femur; (4) in males the characteristic shape of the specialized 
organ on the dorsal surface of the second abdominal segment and of 
the subgenital plate (fig. 10, e, h.). 

Relatively few Nearctic roaches are as large as L, maderae., and none 
of them is likely to be confused with it. Blaherus craniifer Burmeister, 
illustrated by Hebard (1917, pi. 8, fig. 6), is the largest roach in the 
United States, being much larger than maderae. The species of 
Perijylaneta are quite differently marked, and have numerous heavy 
spines on the posterior margins of the femora. Both Eurycotis 
floridana (Walker), of the Southeast, and Hemihldbera tenehricosa 
Rehn and Hebard,^ of the Florida Keys, have short, lobate tegmina 
and wings which are vestigial or lacking. 

A relative of L. maderae recently established in Florida is 
Nauphoeta cinerea (Olivier). It superficially resembles maderae.^ 
though it is smaller, its average length being about 23 millimeters. 
The femora are proportionately stouter and the tegmina scarcely ex- 
tend beyond the apex of the abdomen, while the posterior margin of 
the pronotum is transversely subtruncate, a median projection being 
scarcely evident. Zimmerman (1948, p. 94) and Rehn (1945, p. 274) 
have given photographs of cinerea. This species is apparently a na- 
tive of East Africa now widely distributed in the Tropics by com- 
merce and often intercepted at United States ports. Its potential 
importance is suggested by Illingworth (1942), who described its oc- 
currence in alarming numbers about the feed room of the poultry 
plant at the University of Hawaii. He also mentioned its occurrence 
in heated buildings in England and Germany, though no records of 
permanent establishment in those countries have come to my attention. 

In early December 1951 material of cinerea was received from Jo- 
seph Gross of Tampa, Fla., who stated that it was widely and thor- 
oughly established about Tampa, especially in feed mills. Later, in 
August 1952, Mr. Gross contributed a series of 34 adults and 32 nymphs 

s IlemiMaiera tenebricosa was described from the West Indies, including Nassau in the 
Bahamas, by Rehn and Hebard (1927, p. 271). Caudell (1931) recorded an adult pair 
collected at Key Largo. Fla., by E. A. Popenoe in 1896. Caudell's unpublished notes dis- 
close subsequent discussions of this record with Mr. Popenoe's son, who explained that his 
father was collecting insects primarily for Kansas State College during the 1896 trip and 
that in view of conditions at Key Largo at that time this roach was probably an established 
species rather than an intercepted adventive. A number of roaches are now known to 
be native to the northern Bahamas and extreme tropical Florida, so the occurrence of 
tenehricosa in the Florida Keys Is not illogical. 



COCKROACHES NEW TO UNITED STATES — GURNEY 



47 




Figure 10. — Structural details of Ectobius livens (Turton) and Leucophaea maderae (Fab- 
ricius). a-d, E. livens: a. Dorsal view of apical half of cleared male abdomen; Z», wing; 
c, ventral view of subgenital plate and associated cerci, male; d, apical portion of genital 
hook, male, e-h, L. maderae: e, Dorsal view of second abdominal segment, male, cleared 
and in alcohol;/, dorsal view of apical half of cleared male abdomen; g, genital hook of 
male, enlarged; h, ventral view of subgenital plate and associated structures, male, dry 
preparation. Drawn from specimens as follows: a-d, from Massachusetts specimen; e, 
from Puerto Rico specimen; / and g, from specimen collected in a Tampa, Fla., warehouse; 
A, from New York City specimen. (A-9, apodemes of sternum 9; ca, oblique carinae of 
glandular specialization; gh, genital hook; gld, glandular depression; itr, intercalated 
triangle; me, mesal extension; st, stylus; sap, supra-anal plate; sgp, subgenital plate.) 
228354—53 2 



48 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

to the U. S. National Museum. I inquired especially as to the apparent 
permanence of the infestation, and on June 24, 1952, a colleague of Mr. 
Gross', W. B. Gresham, Jr., replied, "It seems evident to us that 
Nauphoeta is in Florida to stay. They appear to be well adapted and 
quite numerous in the locations where we have noted them. All in- 
festations noted have been within commercial buildings engaged in the 
manufacture of animal feeds. I myself have not noted any out of 
doors." Notes on the Florida infestations by Ratcliffe (1952) and 
Gresham (1952) have appeared. 

Description. — Adults : General form as in plate 2, figure 1 ; tegmina 
and wings extending beyond apex of abdomen about one-fifth their 
length ; width between eyes on vertex about two-thirds length of first 
antennal segment ; a shallow, scarcely wrinkled transverse depression 
in interocular area where eyes are closest ; ventroposterior margin of 
middle and hind femur each with short, apical spine; conspicuous 
spines on posterior femoral margins otherwise lacking; ventropos- 
terior margin of front femur with continuous row of delicate, seta- 
like spines, these slightly shorter toward apex of femur and a few 
noticeably longer ones near base ; front tibia with two elongate groups 
of setae along margins; pulvillus on each tarsal segment, elongate 
on segments 1 and 5; claws equal, unarmed ventrally, with sparse 
setae dorsally ; pronotum obtusely angulate laterally, posterior margin 
with rounded median production; tegmen with cubital veins (dis- 
coidal sectors) oblique; wing (see J. W. H. Eehn, 1951, pi. 9, fig. 102) 
with well-developed axillary and cubital fields, intercalated triangle 
lacking. Dorsal surface of abdomen simple in female, specialized in 
male; second tergum of male (fig. 10, e) with median glandular de- 
velopment consisting of paired oblique carinae (ca) and a brief longi- 
tudinal carina centered on a trifid papilla which largely covers a 
small glandular depression (gld) ; supra-anal plate transverse, with 
median emargination of posterior margin, general shape more quadrate 
when dry (fig. 10, h) than when preserved in alcohol (fig. 10, /) ; sub- 
genital plate unspecialized in female, slightly asymmetrical in male, 
with recurved lateral hooks near styli ; cleared male abdomen with 
apodemes of sternum 9 (subgenital plate) (A9) united anteriorly, 
genital hook (gh) and additional sclerotized elongate rod associated 
with an irregular genital mass; genital hook (fig. 10, g) enclosed in a 
membranous sheath. 

General color pale brown, the tegmen and pronotum marked with 
dark brown as illustrated (pi. 2, fig. 1) ; remainder of body and ap- 
pendages largely dark brown ; clypeus and most of labrum pale ; ab- 
dominal sterna with oblique submarginal marks of brownish-black; 
terga with more heavily pigmented areas as shown (fig. 10, /). Base 
of genital hook with pigmented transverse lines, the apical portion 
much darker. 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 103 PLATE 2 




1, 2, Leucophaea maderae (Fabricius); 1, Adult female, collected in Manila, Philippine 
Islands (length, 47 mm.): 2, last nymphal stage, intercepted at Washington, D.C. 

3-6, Ectohius livens (Turton), all from Falmouth, Massachusetts: 3, 4, Nymphal stages; 
5, cotheca; 6, adult female (length, 9 mm.). 



COCKROACHES NEW TO UNITED STATES — GURNEY 49 

Measurements (in millimeters) : Overall body length, including 
folded tegmina, female 45 to 53, male 38 to 45 ; pronotum, female 9 to 
10.3, male 7.5 to 8.5 ; tegmen, female 37 to 46, male 32 to 38.5. Width 
of pronotum, female 13.3 to 16, male 11.5 to 14. 

Nymphs: Nymphs (pi. 2, fig. 2) ranging from 7.5 to 32 millimeters 
in body length have been examined. The dorsal surface of the thorax 
and abdomen is sharply rugose, with short, microscopic spines, these 
especially conspicuous along the posterior margins of segments; 
much smaller and less evident spines on ventral surface of abdomen. 
General color reddish brown ; each tergum with a submarginal dark 
spot near the base, the basal half of the exposed lateral margin black- 
ish and the apical third yellowish. 

Distnhution. — In the United States Leucophaea maderae is known 
to be established only in the Harlem section of New York City.*' Many 
times each year inspections made at shipping centers detect the species 
with plant products and other articles coming both from the American 
Tropics and the Old World. The distribution of maderae in warm 
countries is very wide, but somewhat irregular, depending upon the 
vagaries of commerce. The specific and the common names are de- 
rived from the fact that this roach was first described from Madeira, 
a small island now belonging to Portugal and located in the Atlantic 
Ocean about 400 miles west of Morocco. Eehn ( 1937, pp. 56-58 ; 1945, 
p. 273) has reasoned (by a detailed analysis of the present occurrence 
of Tnaderae and its congeners) that West Africa is the native home, 
from which commerce (in early centuries — that of the slave trade in 
particular) has carried it to Madeira, the West Indies, Brazil, and 
elsewhere. Ecuador, Colombia, Venezuela, the Guianas, Brazil, and 
Argentina are known to have established colonies of maderae^ as well 
as Panama, Costa Rica, and most of the West Indian islands. Old 
World distribution outside of Africa is more spotty and, to my knowl- 
edge, the species is not yet recorded from India or Australia — even 
though it is known from Java, the Philippines, Hawaii, and Fiji. 
Doubtless, 7naderae now breeds in manj^^ coastal areas from which 
there are no records and it will continue to spread as conditions permit. 

Biology. — In New York City Leucophaea maderae has been "found 
in some abundance in the basement of an apartment of the Harlem 
section," and an observer also writes ^ "From their abundance and the 
presence of all stages of the insect .... I would conclude that they 



•Davis (1940) reported at a meeting held December 15, 1938, that a specimen of 
L. maderae taicen in the Bird House of Bronx Park, New York City, was the first record 
of tliis species in New York State. In the absence of further information, there is a strong 
likeliliood that the specimen in question was an escaped adventive. 

' Specimens from the described infestation were submitted by Ralph E. Heal. Technical 
Director of the National Pest Control Association. Grateful acknowledgment is made of 
Dr. Heal's cooperation in obtaining information on the Harlem Infestation. 



50 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

are breeding in this house." It was also noted that the Harlem roaches 
appeared rather sluggish in their movements. 

In the absence of biological studies of L. maderae in this country it 
is instructive to review the observations of life history and habits 
that have been made elsewhere, particularly the papers of Illingworth 
(1915), Sein (1923), and Pessoa and Correa (1928). 

As is true of species of PeHplaneta^ growth in the Madeira roach 
is slow. The first molt may occur 1 to 4 months after birth. Pinned 
nymphs reared by Illingworth in Hawaii and now deposited in the 
U. S. National Museum suggest that there are six nymphal stages, 
though the number is likely to vary, and in cultures it may sometimes 
be difficult to observe because exuviae reportedly are eaten immedi- 
ately after each molt. Pessoa and Correa report only four "meta- 
morphoses," but that number of molts is almost certainly too small 
for a normal growth cycle. The body length in millimeters of speci- 
mens of various ages collected by Illingworth is as follows: 1 day, 
7.5; 1 month, 7.5; 4 months, 16; 5 months, 18.5; 6 months, 24.5; 7 
months, 29 ; 8 months, 30.7 ; 9 months, 32. Food and environmental 
conditions are doubtless important influences on growth. Sein notes 
maturity being attained in 220 days, but that a year is often required, 
while Pessoa and Correa indicate that adults are obtained in a year, 
with 16 to 18 months the longest observed time. In Sein's cultures 
the first nymphs appeared 4 months after their parents matured. 
Inmiediately following birth, nymphs usually hide beneath the 
mother during the day. According to Pessoa and Correa, this makes 
the adult restless and active in contrast to its usual slow gait. 

In a general paper emphasizing the adaptability and value of 
Leucophaea Trmderae as an experimental animal, Scharrer (1951) 
comments that it thrives on a diet of apples, carrots, and dogfood. 
She reports that there are 30 to 35 young every 3 months, that there 
is an average of 8 molts, and that life expectancy is up to 2V^ years. 
Dr. Scharrer has observed that tumors which resemble malignant 
cancer of higher animals develop in various parts of the roach body 
following the removal of the corpora allata and the corpora cardiaca. 
That such an injury to the nervous system has produced tumors has 
very interesting and practical implications for further research. 
The name "woodroach" that Dr. Scharrer applied to maderae is a 
questionable choice, since the name is most often used for native 
Nearctic species of Parcohlatta. 

Leucophaea is one of several roach genera which are viviparous; 
that is, the eggs are enclosed in a delicate membranous sac which nor- 
mally ruptures before extrusion from the mother's body, or immedi- 
ately thereafter, so that, in effect, the young are born alive. This is 
in contrast to the habit of most roaches which form a heavily-sclero- 
tized, dark-colored capsule or ootheca, usually of characteristic shape, 



COCKROACHES NEW TO UNITED STATES — GURNEY 51 

which may be carried protruding from the body for some time prior 
to hatching, or be deposited loosely or surrounded with a matrix. 

The viviparity exhibited generally takes a special form known as 
ovoviviparity. This occurs in several well-known genera, especially 
Panchlora^ Pycnoscelus^ Nauphoeta^ and Leucophaea^ and Chopard 
(1938, p. 218) notes six subfamilies of roaches (according to the long- 
used classification) in which viviparity occurs. J. W. H. Rehn (1951)' 
has recently proposed a new classification, based on wings, and it is 
significant that many of the viviparous genera fall in what he terms 
the epilamproid complex. In addition to Chopard, Shelf ord (1907), 
Karny (1924, pp. 3-10), and Rau (1941) have discussed viviparity 
among roaches. Hagan (1941; 1951) has described the female re- 
productive system of the viviparous Diploptera^ and Chopard (1950), 
that of Gromphadorhina. 

lUingworth and Sein both recognized that Leucophaea maderae is 
viviparous, but it appears that Pessoa and Correa observed unusual 
or abnormal instances of the Qgg sac being deposited, as they wrote of 
a capsule being placed in the darkest corner of a rearing box and the 
first young appearing 20 days later. In the following paragraphs I 
have attempted to explain this apparent lack of agreement. 

Several preserved adult females of L. maderae received for identifi- 
cation during recent years have exhibited an elongate sac about 20 
millimeters long protruding from the end of the abdomen. In cer- 
tain cases the eggs (varying up to about 40 in number), directed 
transversely with respect to the mother's body and arranged in two 
rows, were undeveloped, but in others nymphs ready for hatching were 
visible — in fact, rupturing of the sac had sometimes occurred. Rehn 
(1937, p. 62) has described his experience in the Belgian Congo with 
the related L. grandis (Saussure) . A female confined in a bottle gave 
birth to 20 living young. The number of young of tnaderae produced 
at one time, according to published records, ranges from 25 to 32. 

The related Pycnoscelus surinamensis (Linnaeus) is known to be 
viviparous, but the literature concerning its habits suggests a lack of 
uniformity in birth or hatching. Watson (1929, p. 58) commented on 
an egg capsule being deposited but the eggs failing to hatch. Caudell 
(1925) also maintained cultures of P. surinamensis, and mentioned 
young born alive and oothecae seldom if ever protruding from the 
female abdomen. In a somewhat fuller account, Zappe (1918) gave 
the opinion that young of surinamensis are either born alive or hatch 
from eggs within 24 hours. Eggs were often laid in soil, but were not 
observed to hatch. When females were injured or excited, they often 
deposited poorly developed egg masses that did not hatch. Similarly, 
lUingworth (1942) reported that the eggs of Nauphoeta cinerea were 
usually kept in the body until living young appeared, but that confined 
individuals sometimes produced aborted egg masses that were ex- 



52 PROCEEDINGS OF THE NATIONAL MUSEUM vol.103 

truded but which dried up without hatching when the delicate enclos- 
ing membrane was exposed to the air. These experiences suggest that 
some of the protruding eggs of L. niaderae that I have noted are the 
result of handling when the specimens were collected and that occa- 
sional Qgg masses may be deposited under certain conditions by this 
ordinarily viviparous species. 

In his important recent paper, Chopard (1950) has discussed the 
anatomy and development of Gromfhadorhina laevigata Saussure 
and Zehntner, of Madagascar. He observed, in the course of rearings 
conducted in Paris, that the female of this viviparous species extrudes 
the soft ootheca nearly to its full extent, then it is drawn into an 
incubating pouch where the eggs undergo incubation for approxi- 
mately 70 days prior to the appearance of the young. Occasionally 
the ootheca is completely extruded, with the result that desiccation 
occurs and no hatching takes place. I am indebted to my colleague 
R. E. Snodgrassfor pointing out the significance of Chopard's obser- 
vations. They explain how the ootheca is transferred from the uterus 
to the incubating chamber in the case of Gromphadorhina. Possibly 
the same habit occurs in certain other viviparous genera, and some 
protruding oothecae that have been seen may represent a stage in this 
normal act. In discussing the transfer of the developing eggs in 
the viviparous Difloptera^ Hagan (1951, p. 299) states that the 
oocytes pass from the ovarioles to the lower end of the common ovi- 
duct, where "they are directed by the ovipositor from the genital 
chamber ventrally into the open end of the uterus." 

In Brazil, mating of Leucofhaea maderae occurs mainly during the 
warm and rainy season, according to Pessoa and Correa, who say that 
copulation may occupy 20 to 30 minutes and takes place with the pair 
end to end facing in opposite directions. Prior to mating, the female 
is described as opening her wings and drawing them along the ground, 
at the same time producing a sound by vibrating them. The scent 
gland on the dorsum of the male presumably is attractive to the female 
at mating time, but its function is not definitely known. 

lUingworth stated that when disturbed the Madeira roach stridu- 
lates very noticeably, and he believed the sound is produced by rubbing 
the posterior margin of the pronotum over the mesonotum. Accord- 
ing to Chopard (1938, p. 286), several roaches stridulate delicately by 
rubbing the border of the pronotum upon the mesonotum or upon the 
strongly denticulate base of the costal vein of the tegmen. To test the 
possibilities of stridulation, I have relaxed dry specimens of both sexes 
of L. maderae^ then manipulated the body parts with my fingers. A 
low, squeaking sound is consistently obtained in either sex by rubbing 
the lateroposterior margin of the pronotum on the basal costal margin 
of the tegmen. Both surfaces are heavily sclerotized, turned to oppose 



COCKROACHES NEW TO UNITED STATES — GURNEY 53 

each other, nonpubescent, and are finely rugose so as to make stridula- 
tion possible. 

In tropical regions where it is established, Z. maderae is definitely a 
domiciliary species, though, like certain other roaches often associated 
with man, it is capable of living apart from him in a purely wild state. 
From available records, however, it is not clear how frequently it has 
been collected entirely unassociated with man-made surroundings. 
According to Alfken (1904, p. 565) more than 50 years ago a German 
collector, H. Schauinsland, in 1896 or 1897 found mfiderae in native 
huts on the Hawaiian Island of Molokai, commerce already having 
carried it to certain Pacific areas. However, in spite of this long estab- 
lishment in the Hawaiian group, no mention of its occurrence in fields 
is made by Williams ( 1931 ) . In July 1950, in the Venezuelan State of 
Aragua, Dr. Ernst Schwarz ^ collected it coming to lights in fair num- 
bers, but whether it was breeding in the adjacent forest or associated 
with nearby habitations is not known. Likewise, specimens collected 
by the late C. F. Baker in the Philippines before 1927 bear the collect- 
ing label "Mt. Maquiling," and they may have bred under entirely nat- 
ural surroundings. However, as I saw during my visit to Mt. Maquil- 
ing in 1945, at the base and on the lower slopes there are ample oppor- 
tunities for this roach to remain associated with man. The species 
frequently flies actively and has often been taken at lights, including 
those on porches, in field camps, or of automobiles. On the other hand, 
Sein states that Leucophaea maderae does not fly much when indoors 
at night as part of an infestation. L. puerilis Rehn, considered its 
closest relative, is a strictly endemic, forest type in West Africa. 

Source of introduction. — The New York City infestation is reported 
to be localized in apartment buildings occupied by people from Puerto 
Rico and, since there has been a good deal of movement — much of it 
by air — from that island in the past few years, there is little doubt that 
Puerto Rico is the source of the infestation. 

Probable importance. — In Puerto Rico the Madeira roach most often 
occurs in fruit stores and markets. It is especially fond of grapes. It 
is considered very gregarious and develops large, localized colonies. 
Wolcott (1950, p. 43) records about a bushel having been swept from 
one store. Warehouses and other buildings are often infested. When 
handled or otherwise disturbed, Leucophaea maderae produces an odor 
described as especially offensive. Houses infested by the species usu- 
ally do not contain Periplaneta or Blattella. There is a strong possi- 
bility that maderae., if allowed to spread, will gradually develop into a 
serious pest in our larger cities. In the light of its wide occurrence in 
the Bahamas and other West Indian islands, tropical Florida would 
probably be a suitable habitat, either outdoors or in buildings without 
central heating. 

« Associated with the Venezuela Plague Mission (Commander J. M. Amberson, D.S.N., 
Dr. Ernst Schwarz, and Mrs. Schwarz). 



54 PROCEEDESTGS OF THE NATIONAL MUSEUM vol.108 

References 

Alfken, Johann D. 

1904. Beitrag zur Insectenfauna der Hawaiischen und Neuseelandischen 
Inseln. Zool. Jahrb., System., vol, 19, pp. 561-628, 1 pi. 
Back, Ebnest A. 

1937. The increasing Importance of tlie cockroach, Supella supellectilium 

Serv., as a pest in the United States. Proc. Ent. Soc. Washington, 
vol. 39, pp. 205-213, 2 pis., 2 figs. 
Blaib, Kenneth G. 

1934. A note on the British species of EctoUus Steph. Ent. Monthly Mag., 

vol. 70, pp. 157-159. 

1935. [British species of EctoUus. '[ Proc. South London Ent. Nat. Hist. 

Soc, 1934-35, p. 5. 
Brown, E. B. 

1952. Observations of the life-history of the cockroach EctoMus panzeri 
Stephens ( Orth., Blattidae ) . Ent. Monthly Mag., vol. 88, pp. 209-212. 
Beunneb de Wattenwyl, Chakles 

I860. Nouveau syst^me des Blattaires. 426 pp., 13 pis., Vienna. 
BuBE, Malcolm 

1936. British grasshoppers and their allies. 164 pp., 6 pis., 56 figs., 40 maps, 

London. 
Caudell, Andrevv^ N. 

1925. Pycnoscelus surinamensis Linnaeus (Orthoptera) ; on its nymphs and 
the damage it does to rose bushes. Proc. Ent. Soc. Washington, vol. 
27, pp. 154-157, 2 figs. 
1931. Notes on Blattidae, adventive to the United States. Ent. News, voL 
42, p. 204. 
Chopard, Lucien 

1938. La biologie des Orthopt&res. Encyclopedic entomologique, ser. A, vol. 

20, pp. 1-541, 4 pis., 453 figs. Paris. 
1943. Orthoptferoides de I'Afrique du Nord. Faune Empire Fran^ais, vol. 1, 

pp. 1-450, 658 figs., Paris. 
1947. Atlas des AptSrygotes et Orthopt^roides de France. Nouvel atlas 

d'entomologie, vol. 2, pp. 1-111, 12 pis., 12 figs. Paris. 

1950. Sur I'anatomie et le developpement d'une blatte vivipare. Proc. 8th 

Int. Cong. Ent., Stockholm, 1948, pp. 218-222, 6 figs. 
Davis, William T. 

1940. [Leucophaea recorded.] Bull. Brooklyn Ent. Soc, vol. 35, p. 35. 
Flint, Oliver S. 

1951. A new cockroach record from the United States. Bull. Brooklyn Ent. 

Soc, vol. 46, p. 53. 
Gaunitz, C. B. 

1935. Till frSgan om EctoMus lapponictis L. formente skadegSrelse inomhus 

i Lappland. Ent. Tidskr., vol. 56, pp. 138-150. 

1936. EctoMus lapponicus L. als Vorratsschadling in Lappland, eine alte 

sicher unrichtige Vermutung in neuer Beleuchtung. Konowia, vol. 

15, pp. 162-166. 
Gould, George E., and Deat, H. O. 

1940. The biology of six species of cockroaches which inhabit buildings. 

Purdue Agr. Exp. Stat. Bull. 451, pp. 1-31, 13 figs. 
Gresham, William B. 

1952. [Note on 2VaM2)/ioe/a in Florida.] Florida Ent, voL 35, p. 77. 



COCKKOACHES NEW TO UNITED STATES — GURNEY 55 

Hagan, Haeold R. 

1941. The general morphology of the female reproductive system of a vivip- 

arous roach, Diploptera dytiscoides. Psyche, vol. 48, pp. 1-9, 1 pi., 

2 figs. 
1951. Embryology of the viviparous insects. 472 pp., 160 figs., New York. 
Hebard, Morgan 

1917. The Blattidae of North America north of the Mexican boundary. 

Mem. Amer. Eut. Soc, No. 2, pp. 1-284, 10 pis., 1 fig. 
1943. Australian Blattidae of the subfamilies Chorisoneurinae and Ecto- 

biinae. Monogr. Acad. Nat. Sci. Philadelphia, No. 4, pp. 1-129, 

14 pis. 
Illingw^obth, James F. 

1915. Notes on Hawaiian roaches. Proc. Ent. Soc. Hawaii, vol. 3, pp. 136-140. 

1942. An outbreak of cockroaches, Nauphocta cinerea (Olivier), in Hawaii. 

Proc. Ent. Soc. Hawaii, vol. 11, pp. 169-170. 
Kabny, Heinkich H. 

1924. Beitrage zur malayischen Orthopterenfauna. Treubia, vol. 5, pp. 1-234, 
1 pi., 85 figs. 
Kbauss, Hermann 

1892, Systematische Verzeichnis der Canarischen Dermapteren und Orthop- 
teren mit Diagnosen der neuen Gattungen und Arten. Zool. Anz,, 
vol. 15, pp. 163-171. 
Lucas, William J. 

1920. A monograph of the British Orthoptera. Ray Soc, Monogr. 104, 264 

pp., 25 pis., 25 figs., London. 
1928. Notes on British Orthoptera, including Dermaptera, in 1927. Ento- 
mologist, vol. 61, pp. 78-81. 
Pessoa, Samuel B., and Correa, Clovis 

1928. Notas sobre a biologie de Rhyparohia maderae, Fabr. Rev. Biol. Hyg., 
Sao Paulo, vol. 1, pp. 83-87, 1 pi., 3 figs. 
Princis, K. 

1936. Erganzungen und Berichtigungen zur Orthopterenfauna Lettlands, I. 

Folia Zool. Hydrobiol., vol. 9, pp. 90-92. 
Ramme, Willy 

1923. Vorarbeiten zu einer Monographie des Blattidengenus EctoMus Steph. 

Arch. Naturg., Berlin, ser. A, vol. 89, pp. 97-145, 2 pis, 55 figs. 
1949. Ein neuer EctoMus vor Sizilien (Blatt.) Opuscula Ent., vol. 14, p. 158, 

1 fig. 

1951. Zur systematik faunistik und biologie der Orthopteren von Siidost- 

Europa und Vorderasien. Mitt. Zool. Mus. Berlin, vol. 27, pp. 1^31, 
39 pis., 134 figs. 
Ratcliffe, .Tack 

1952. Lobster roach. Pest Control, vol. 20, pp. 44, 54. 
Rau, Phil 

1941. Cockroaches: The forerunners of termites. Ent. News, vol. 52, pp. 
256-259. 
Rehn, James A. G. 

1903. Studies in American Blattidae. Trans. Amer. Ent. Soc, vol. 29, pp. 
259-290. 

1931. African and Malagasy Blattidae, Part I. Proc. Acad. Nat. Sci. Phila- 
delphia, vol. 83, pp. 305-387, 5 pis. 

1937. African and Malagasy Blattidae, Part III. Proc. Acad. Nat. Sci. 

Philadelphia, vol. 89, pp. 17-123, 4 pis. 



56 PROCEEDENGS OF THE NATIONAL MUSEUM vol. loa 

Rehn, James A. G. — Continued 

1945. Man's uninvited fellow traveler — the cockroach. Sci. Monthly, voL 61, 
pp. 265-276, 11 figs. 
Rehn, James A. G., and Hebakd, Morgan 

1927. The Orthoptera of the West Indies, No. 1. Blattidae. Bull. Amer. 
Mus. Nat. Hist., vol. 54 (Art. 1), pp. 1-320, 25 pis. 
Rehn, John W. H. 

1950. A key to the genera of North American Blattaria, including estab- 

lished adventives. Ent. News. vol. 61, pp. 64-67. 

1951. Classification of the Blattaria as indicated by their wings. Mem. 

Amer. Ent. Soc. No. 14, pp. 1-134, 13 pis., 5 figs. 
Schabkee, Berta 

1951. The woodroach. Sci. American, vol. 185, no. 6, pp. 59-62, 5 figs. 
Seabba, a. F. de 

1942. Contribuicoes par o inventario da fauna lusitanica. Insecta Orthop- 
tera. Mem. Mus. 2k)ol. Univ. Coimbra, Portugal, vol. 127, pp. 1-26. 
Sein, Francisco 

1923, Cucarachas. Puerto Rico Estac. Exp. Insul., Circ. 64, pp. 1-12, 9 figs. 
Shelford, Robert W. O. 

1907. Viviparity amongst the Blattidae. Trans. Ent. Soc. London, 1906, pp. 
509-514. 
Watson, John R. 

1929. [Rearing of Pycnoscelus.} Univ. Florida Agr. Exp. Stat., Ann. Rep. 
1929, p. 58. 
Williams, Francis X. 

1931. Handbook of the insects and other invertebrates of Hawaiian sugar 
cane fields. 400 pp., 41 pis., 190 figs., Honolulu. 
WoLCOTT, George N. 

1950. The insects of Puerto Rico. Journ. Agric, Univ. Puerto Rico, vol. 32 
(1948), pp. 1-224, illus. 
Zappe, Max P. 

1918. A cockroach pest of greenhouses. Connecticut Exp. Stat. Bull. 203, 
pp. 802-313. 
Zimmerman, Elwood C. 

1948. Insects of Hawaii, vol. 2, pp. 1^75, 228 figs., Honolulu 



U. S. GOVERNMENT PRfNTINS OFFICE: 1993 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 




SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 



Vol. 103 Washington : 1953 No. 3316 

BITING MIDGES OF THE HELEID GENUS STILOBEZZIA IN 

NORTH AMERICA 



By Willis W. Wirth ' 



Siilobezzia Kieffer, 1911, is a rather large, widespread genus of 
predaceous midges which has reached its greatest development in 
the warmer parts of the New World. For North America 11 species 
are listed in Johannsen's recent list (1943b); this total is now raised 
to 18 by the present new records and species, and no doubt many 
more await discovery. The species of Stilobezzia are rather easily 
characterized, and all the Nearctic species known to Johannsen were 
correctly placed by him, with the exception of S. uncinata Johannsen, 
1943, which I believe to represent the female of Parabezzia petiolata 
Malloch, 1915. 

The types of the species described here are in the U. S. National 
Museum, the collections of which furnished the bulk of the material 
studied. I am indebted to Dr. Henry K. Townes, of North Carolina 
State College, for the loan of specimens from his collection, and to 
Mrs. Elisabeth C. Beck and J. A. Mulrennan, of the Florida State 
Department of Health, for their kindness in sending a large series of 
light trap collections from which I sorted most of the Florida specimens. 

Family Heleidae 

Genus Stilobezzia Kieflfer, 1911 

Diagnosis. — Body rather slender and nearly bare. Eyes bare. 
Female antenna with segments 3-10 oval, 11-15 long and cylindrical; 



' Bureau of Entomology and Plant Quarantine, Agricultural Research Administration, U. S. Depart' 
ment of Agriculture. 

232991—53 1 57 



58 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

male antenna with well-developed plumes. Mesonotum usually with 
few or no long bristles; humeral pits present but usually inconspicuous. 
All legs slender; femora without spines or (in Eukraiohelea) with a 
few on forelegs; basitarsus and fifth segment with or without ventral 
spines; fourth segment cordiform, fifth not enlarged; female claws 
large and very unequal or a single long claw with basal barb on each 
leg, male claws small and equal; empodium absent. Wing rather 
long; microtrichiae present; macro trichiae present toward tip (in 
Neostilobezzia) or absent; costa extending to two-thirds wing length 
or more; usually two radial cells, second much larger than first, 
which may be entirely absent (in Eukraiohelea) or greatly reduced; 
intercalary fork distinct; crossvein r-m more or less oblique; median 
fork with long petiole, M2 not interrupted at base; anal vein straight, 
not interrupted in middle. Male genitalia with basistyles simple or 
with lobe on inner margins ; dististyles long ; ninth sternite very short ; 
ninth tergite usually rounded caudad with setigerous apicolateral 
lobes; aedeagus reduced to a pair of oblique lateral sclerites; parameres 
a pair of large, highly sclerotized, greatly modified, submedian proc- 
esses with flaring basal apodemes. 

Keys to the Nearctic species of Stilobezzia 

Females 

1. Only one long radial cell present; fore femur with 2 or 3 ventral spines 

(subgenus Eukraiohelea) 1. elegantula (Johannsen) 

Two radial cells present ; fore femur unarmed 2 

2(1). Wing with macrotrichiae at tip (subgenus Neostilobezzia) 3 

Wing entirely bare (subgenus Stilobezzia) 5 

3 (2). Body hairs very strong and erect; wings hyaline; legs, scutellum, and 

halteres yellow 3. lutea (Malloch) 

Body hairs soft and decumbent; midlegs and hind legs brown 4 

4(3). Wing infuscated brown; scutellum and halteres yellowish. 

4. stonci, new species 
Wing hyaline; scutellum and halteres brownish 2. fuscula Wirth 

5 (2). Wing with 2 or 3 dark spots on anterior margin 6 

Wing hyaline, unmarked 9 

6 (5). Shining yellow; mesonotum polished yellow, with pair of longitudinal 

brown bands, anterior spine absent; legs yellow, narrow apices of hind 
femur and tibia brown; wing hyaline, with brown spots at crossvein 

and apex of second radial cell 9. coquilletti Kieffer 

. Mesonotum bro^yn with silvery, pruinose areas and prominent anterior 
spine; legs extensively brown banded or punctate; wing brownish, 
anterior margin with 2 or 3 dark spots 7 

7 (6). Femora and tibiae extensively yellow, with small brown spots at bases of 

hairs; wing hyaline, with two brown spots; abdomen mostly yellowish 

above 14. punctipes, new species 

Femora and tibiae brown with yellowish rings; wing infuscated, with 2 
or 3 brown spots and a subapical pale area; abdomen black above with 
narrow, distal, silvery pruinose, segmental bands 8 



HELEID MIDGES, GENUS STILOBEZZIA — WIRTH 59 

8 (7) . Three dark wing spots including one at apex of first radial cell; fifth tarsal 

segment with ventral batonnets 15. rabelloi Lane 

Two dark wing spots, none at apex of first radial cell; fifth tarsal segment 
without ventral batonnets 6. beckae, new species 

9 (5) . Mesonotum entirely polished black 10 

Mesonotum yellow, green, or brown, not entirely black 13 

10 (9). Abdomen pohshed black 11 

Abdomen green 12 

11 (10). Midfemora and hind femora black except at extreme bases. 

5. antennalis (Coquillett) 
Legs uniformly yellow 7. bicolor Lane 

12 (10). Fifth tarsal segment with ventral batonnets. _ 10. diversa (Coquillett) 

Fifth tarsal segment without ventral batonnets. 

12. pallidiventris (Malloch) 

13 (9). Shining green; third or fourth abdominal tergites extensively black. 14 

Yellow or brown, not green; abdomen without black bands or spots. 15 

14 (13). Mesonotum uniformly pale green; distal fourth of hind femur black. 

18. viridis (Coquillett) 

Mesonotum largely dark brown; hind femur with only spot at apex 

black 11. glauca Macfie 

15 (13). Mesonotum shining brown; wing with a beadlike swelling near apex of 

second radial cell 16. sybleae, new species 

Mesonotum pruinose brown; wing venation normal 16 

16 (15). Mesonotum dark brown, with pattern of large pruinose gray spots; 

scutellum dark; tibiae with sub-basal pale rings.. 13. pruinosa Wirth 
Mesonotum uniform light grayish brown; tibiae unbanded 17 

17 (16). Scutellum and femora yellow 8. bulla Thomsen 

Scutellum and femora brown 17. thomscnae, new species 

Male genitalia 

1. Lateral sclerites of aedeagus greatly reduced or absent, instead a trans- 
verse sclerite between inner margins of basistyles beyond base 2 

Lateral sclerites of aedeagus well developed and oblique, with bases at 
inner ventral margins of basistyles 3 

2 (1). A large rounded lobe at distal third of inner margins of basistyles; 

transverse sclerite with pointed, curved ends; apicolateral lobes of 

ninth tergite widely spaced and triangular 8. bulla Thomsen 

Basistyles simple; transverse sclerite with blunt ends; apicolateral lobes 
fingerlike and close together 17. thomsenac, new species 

3 (1). Lateral sclerites of aedeagus sinuate, irregularly bent, or apex forked; 

basistyle without lobe, or if one present it bears a strong spine; apico- 
lateral lobes of ninth tergite usually slender and close together 4 

Lateral sclerites of aedeagus straight, at least on distal half; basistyle 
with mesal lobe; apicolateral lobes usually low and rounded 10 

4 (3). Basistyle with a strong spine midway on inner margin, arising from a low 

lobe also bearing a fine hair; aedeagus with strong subapical tooth 
on outer side, appearing forked at end; parameres long and slender 

with long pointed tips 2. fuscula Wirth 

Basistyle simple ; aedeagal sclerites without strong tooth 5 



60 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

5 (4). Parameres greatly expanded from bases, with obliquely truncated 

apices bearing a short lateral tooth and a long, curved, distal tooth. 

4. stonei, new species 
Parameres broadest at bases or without truncated apices 6 

6 (5). Stems of parameres very slender and rodlike 7 

Stems of parameres bladelike 9 

7 (6). Apices of parameres slightly expanded, palplike; aedeagal sclerites 

sinuate, stouter toward apices; ninth sternite with deep, quadrate, 
mesal notch ; dististyles very broad, with truncate apices. 

1. elegantula (Johannsen) 

Apices of parameres sharp-pointed and bent ventrad; aedeagal sclerites 

bent more than twice 8 

8 (7). Membrane of ninth sternite bare; tips of aedeagal sclerites recurved 

hooklike; dististyles very slender 6. beckae new species 

Membrane spiculate; tips of aedeagel sclerites snoutlike and crossing 
each other; dististyles stout 14. punctipes, new species 

9 (6). Parameres with an extra lateral pair of straight, bladelike arms half 

as long as inner arms ; membrane of ninth sternite bare. 

15. rabelloi Lane 

Parameres simple, without lateral arms, expanded subapically, with tips 

abruptly narrowed; membrane spiculate 3. lutea (Malloch) 

10 (3). Parameres with knobbed apices over twice as broad as the very slender, 

stalklike stems; basistyles very large and globular; dististyle very 

slender to base 18. viridis (Coquillett) 

Parameres otherwise, without knobbed apices 11 

11 (10). Parameres with greatly swollen, conical bases and very slender, tapered 

apices 12 

Parameres with stems of subequal diameter or swollen towards 
apices 13 

12 (11). Parameres with basal 0.6 swollen, the pointed apices abruptly bent 

towards base on outer side 10. diversa (Coquillett) 

Parameres with only basal 0.2 swollen, distal portions consisting of very 
slender, curved, saber-shaped blades crossing each other in middle. 

16. sybleae, new species 

13 (11). Parameres nearly straight, with apices abruptly narrowed and bent 

over 14 

Parameres sinuate or with broad, toothlike, lateral, subapical expan- 
sions 15 

14 (13). Parameres with apices bent outward 7. bicolor Lane 

Parameres with apices bent inward 11. glauca Macfie 

15 (13). Lobe of basistyle quadrate, as long as broad; parameres slender and 

markedly sinuate; dististyles greatly curved. 

5. antennalis (Coquillett) 

Lobe of basistyle consisting of only a hump on inner margin ; parameres 

very broad and stout ; dististyle nearly straight 16 

16 (15). Each paramere with broad, toothlike, lateral, subapical expansion about 

half as broad as length of dististyle, with a slender, ventrally curved 
spine from apex on inner margin; dististyle tapered. 

9. coquilletti Kieflfer 

Each paramere swollen slightly midway to apex without subapical lateral 

expansion; dististyle with broad, blunt apex 13. pruinosa Wirth 



HELEID MIDGES, GENUS STILOBEZZIA — WIRTH 61 

Table 1. — Proportions of segments of hind legs of female Stllobezzia species. 



Species 



Cx 


Tr 


F 


Ti 


Ti 


T2 


Ta 


T, 


20 


10 


60 


60 


30 


10 


5 


4 


25 


10 


60 


65 


32 


15 


6 


4 


40 


15 


90 


90 


40 


20 


9 


6 


25 


10 


70 


70 


30 


18 


7 


5 


20 


12 


55 


55 


28 


12 


3 


3 


30 


15 


75 


75 


40 


18 


6 


5 


15 


7 


40 


35 


17 


10 


4 


3 


15 


7 


35 


35 


15 


8 


3 


2 


30 


15 


80 


85 


40 


18 


5 


5 


13 


6 


35 


32 


16 


6 


3 


3 


20 


10 


65 


70 


33 


15 


4 


4 


20 


10 


50 


50 


28 


10 


4 


4 


30 


15 


80 


80 


40 


20 


6 


4 


40 


15 


85 


95 


45 


25 


10 


5 


15 


7 


45 


50 


25 


12 


3 


2 


20 


10 


55 


55 


28 


12 


3 


3 



9. 
10. 
11. 
12. 
13. 
14. 
15. 
16. 
17. 



elegant ula 

fuscula 

lutea 

stonei 

antennalis 

beckae 

bicolor 

bulla 

coquilletti 

diversa 

glauca 

pallidiventrio 

pruinosa 

punctipes 

rabelloi 

sybleae 

viridis 



12 

8 

15 

10 



10 
5 

13 
5 

10 



10 

15 

5 

8 



Subgenus Eukraiohelea Ingram and Macfie, 1921 

Eukraiohelea Ingram and Macfie, Ann. Trop. Med. Parasit., vol. 15, p. 347, 1921. — 
Johannsen, Arch. IlydrobioL, Suppl., vol. 9, p. 430, 1931; Journ. New York 
Ent. Soc, vol. 42, p. 344, 1934.— Macfie, Ann. Trop. Med. Parasit., vol. 34, 
p. 22, 1940.— Tokunaga, Tenthredo, vol. 3, p. 344, 1940.— Johannsen, Ann. 
Ent. Soc. Amer., vol. 36, p. 781, 1943; Bull. Bishop Mus. No. 189, p. 190, 1946. 
(Genotype, Eukraiohelea africana Ingram and Macfie, 1921; designated by 
Macfie, 1940.) 

Diagnosis. — First radial cell absent, the r-m crossvein nearly form- 
ing a straight line with Ri, the latter arising just a little more toward 
base of wing; fore femora with 2 or 3 ventral spines but not swollen; 
male aedeagus with hyaline posterior membrane, parameres long and 
very slender. 

Discussion.' — Johannsen (1934) placed Eukraiohelea as a subgenus 
of Parabezzia Malloch, 1915. However, as shown by a comparison 
of the male genitalia of P. petiolata Malloch, 1915, with those of the 
four species of Eukraiohelea whose males have been described, the 
latter are much more closely related to Stilobezzia. The following 
species have been correctly placed in Eukraiohelea: E. africana Ingram 
and Macfie, 1921, E. versicolor Ingram and Macfie, 1921, and E.foyi 
Ingram and Macfie, 1922, all from West Africa; Stilobezzia (Eukraio- 
helea) aberrans Johannsen, 1931, from Java; S. (E.) esakiana Tokunaga, 
1940, from the Caroline Islands; and Eukraiohelea amnigena Macfie, 
1935, from Brazil. Eukraiohelea inusitata Johannsen, 1946, from 



62 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Guam is apparently a synonym of esakiana Tokunaga. Palpomyia 
dorsofasciata Lutz, 1914, from Brazil was correctly referred to Eukraio- 
helea by Macfie (1935). Parabezzia poikiloptera Ingram and Macfie, 
1922, from West Africa should be referred to Eukraiohelea as tenta- 
tively suggested by Johannsen (1946), but the position of S. ugandae 
Ingram and JMacfie, 1923, from East Africa, which Johannsen also 
believed to belong here, is more doubtful. De Meillon (1938), more- 
over, states that versicolor and poikiloptera should be placed in the 
subgenus Stilobezzia, since material he determined as these two species 
from Lourengo Marques, Mozambique, possessed a small but distinct 
first radial cell. 

1. Stilobezzia (Eukraiohelea) elegantula (Johannsen), 1908, new combination 

Figure II, g 

Bezzia elegantulus Johannsen, Kansas Univ. Sci. Bull., vol. 4, p. 109, 1907 ( 9 , 

Kansas) . 
Probezzia elegantula, Malloch, Proc. Biol. Soc. Washington, vol. 27, p. 137, 1914. 
Parabezzia elegantula, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 359, 

1915. — Johannsen, Journ. New York Ent. Soc, vol. 42, p. 345, 1934. 
Eukraiohelea elegantula, Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943. 

Description. — Female: Length 2.0 mm., wing 1.5 mm. by 0.7 mm. 
A yellowish species, some specimens with a greenish tinge; antennae 
brown, pedicel and narrow bases of flagellar segments yellow; palpi 
dark brown. Mesonotum deep yellow, with broad sublateral brown 
bands; scutellum yellow in center, ends brown; postscutellum brown; 
pleura grayish, a large black spot on mesopleuron. Legs yellow; coxae 
dark below; trochanters brown; apex of hind femur, base of hind tibia, 
apices of all tibiae and narrow apices of all tarsal segments brown. 
Wings, including veins, grayish hyaline ; halteres yellow, sides of knobs 
black, flat end of knobs pale green. Abdomen dark grayish brown, 
apices of tergites with narrow white bands. 

Antennae very long and slender. Mesonotum with long black 
hairs in rows; scutellum with four long black marginal bristles and a 
few short hairs. Legs with hairs fine except on hind tibia where they 
are long and black ; proportions of segments of hind leg as in table 1 ; 
fore femur with two or three small black spines midway on inner 
margin; basitarsus with a long black basal and a smaller distal spine 
on midlegs and hind legs; fifth tarsal segment with two pairs of long 
black ventral batonnets; claws slender and unequal, the outer claw 
as long as fifth segment, the other about a third as long. Wing with- 
out macro trichiae; first radial cell absent, r-m crossvein and Ri nearly 
forming an oblique line, lengths of the two sections of costa in ratio 
of 0:6; petiole of media as long as r-m crossvein. 



HELEID MIDGES, GENUS STILOBEZZIA — ^WIRTH 63 

Male: Ninth sternite nearly four times as broad as long, with deep 
quadrate emargination three-fourths way to base on mesal third, with 
spiculate margms, the posterior membrane bare; ninth tergite rounded, 
with a pair of very small iingerlike setigerous apicolateral lobes. 
Basistyles simple, about twice as long as broad; dististyles about two- 
thirds as long as basistyles, very broad with truncate tips. Aedeagus 
with an oblique pair of rather stout, slightly sinuate, sclerotized bars 
with pointed apices; an irregular hyalme membranous lobe from pos- 
terior margins. Parameres very long, slender and rodlike, with slender 
lateral apodemes; stems about a fourth again as long as basistjdes, 
subparallel, the apices slightly swollen, palplike, and bent ventro- 
laterad. 

Type. — In Snow collection. University of Kansas, 9, Kansas, 
Douglas County, Lawrence. 

Material examined. — Florida: Crystal River, Citrus County, July 
7, 1949, Sept. 18, 1950, Hudson, 2 99, Miami, Dade County, Oct. 15, 
1947, Bm-en, 1 9; Panama City Beach, Bay County, July 2, 1950, 
McElvey, 1 cf 

Louisiana: Baton Rouge, Apr. 30, 1947, Wirth, 1 9. 

Jamaica: Newport, Feb. 22, 1937, Chapin and Blackwelder, 1 9. 

Remarks. — S. (E.) africana Ingram and Macfie, from West Africa, 
the genotype of Eukraiohelea, is almost identical with elegantula, but 
is smaller and the male genitalia differ markedly; the ninth sternite 
does not have the mesal excavation, the dististyles have the apices 
slenderer and rounded, the apices of the parameres are straight and 
pointed, and the membrane of the aedeagus is prolonged mesad in a 
sharp cone. 

Subgenus Neostilobezzia Goetghebuer 

Neostilobezzia Goetghebuer, in Lindner, Die Fliegen der Palaearktischen Region, 
Lief. 78, p. 53, 1934. (Genotype, Ceratopogon ochraceus Winnertz, 1852, 
by present designation.) 

Discussion. — Goetghebuer (1934) distinguished this subgenus from 
Stilohezzia by the presence of macrotrichiae at the end of the wing, 
the wing of Stilohezzia being entirely bare. In addition, all the species 
known to me are yellowish or brown in ground color, varying from 
almost whitish to almost black, and the male genitalia have the 
aedeagal sclerites decidedly bent or humped in the middle and the 
basistyle lacks the inner lobe. Species with these characters are 
widely distributed around the world. It is especially significant that 
most of the species described from the temperate regions belong to 
this subgenus, while the tropical species are predominantly the sub- 
genus Stilohezzia. 



64 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

2. Stilobezzia {Neostilobezzia) fuscula Wirth, 1952 

Figure 11, e 

Stilobezzia fuscula Wirth, Univ. California Publ. Ent., vol. 9, p. 204, 1952 ( 9 , 
California) . 

Diagnostic characters.— This species resembles S. (N). lutea (Mal- 
loch), 1918, in its uniformly yellowish color, unmarked wings with 
macrotrichiae, and mesonotum with indistinctly darker vittae; but 
differs in being somewhat duller and darker, with dark antennal 
pedicel and halteres, and in having the body hairs much softer. 

Description. — Female: Length 1.4 mm., wing 1.5 mm. by 0.5 mm. 
Head grayish brown pruinose, pedicel of antennae and palpi brown, 
antennal flagellum yellowish brown; segments in proportion of 
20 : 12 : 12 : 12 : 12 : 12 : 12 : 12 : 20 : 20 :20 : 20 : 30. Palpal segments m pro- 
portion of 4:8:15:10:10, third segment with three long, stalked sen- 
sillae. 

Mesonotimi dull yellowish brown, with coarse pruinosity, a narrow 
median longitudinal vitta and a broad patch on each side from humeral 
pits to wing base darker brown; most of bristles removed, but those 
few remaining long and dark brown. Scutellum bright yellow in 
middle, brownish on sides, apparently with four marginal bristles. 
Postscutellum and plem'a yellowish brown pruinose. Legs dull dark 
yellowish; hairs short, except row of long bristles on outer edge of hind 
tibia; inner claw as long as fifth tarsal segment. 

Wings grayish hyaline, anterior veins brownish; second anterior 
radial cell about five times as long as fu'st; macrotrichiae thick at apex 
of cell R5 and along and beyond intercalary fork, a few also along tip 
of veins Mi and M2 and at apex of cell Mi. Halteres brown, flat end of 
knob dull yellowish. 

Abdomen dull dark brown, a patch of long brown hairs on sides 
of first segment. Spermathecae two, oval, with ducts sclerotized 
a short distance; a rudimentary third spermatheca present. 

Male: Similar to the female but with the usual sexual differences; 
antennal plumes hght brown; thorax and abdomen infuscated light 
brown; scutellum yellow. 

Ninth sternite about three times as broad as long, with mesal 
posterior excavation halfway to base, posterior membrane bare; ninth 
tergite rather long and rounded, with a pair of long, rounded, seti- 
gerous, apicolateral lobes. Basistyle nearly three times as long as 
broad, with a black spine borne on a small lobe with a long fine hau- 
about midway of mesal margin; dististyle as long as basistyle, slender 
and slightly curved to apex. Aedeagus with a pair of oblique, lateral, 
sclerotized bars with median apices forked, the inner arm in line 
with base and longer than the other arm. Parameres with slender 



HELEID MIDGES, GENUS STILOBEZZIA — WIRTH 65 

basal apodemes, stems long and slender, with bases slightly swollen, 
each with apex gradually narrowed to a fine apical filament abruptly 
bent and recurved on distal fourth. 

Type.—VSNM 59946, 9, California, Tulare County, 6 miles east of 
Orosi. (In the original publication the sex was erroneously given 
as cT.) 

Material examined.— Caliiormsi: 6 miles east of Orosi, Tulare 
County, July 8, 1947, Wirth, 2 99 (type and paratype). 

Utah: Moab, June 8, 1948, Knowlton, Harmston, and Wood, at 
light. Id". 

Remarks. — The male genitalia of S.fuscula are practically identical 
with those of S. macfiei Lane, 1947, from Brazil. However, the latter 
species has the mesonotum polished black, with lighter areas, and 
under certain light a spot on each side before wing base has purplish 
reflections. Moreover, the absence of macrotrichiae on the wings 
would place S. macfiei in the subgenus Stilobezzia. 

3. Stilobezzia {Neostilohezzid) lutea (Malloch), 1918 

Figure 11, 6 

Hartomyia gilva Malloch, not Coquillett (misidentification) , Bull. Illinois State 

Lab. Nat. Hist., vol. 10, p. 343, 1915. 
Hartomyia lutea Malloch, Bull. Brooklyn Ent. Soc, vol. 13, p. 18, 1918, ( 9 , Illinois). 
Stilobezzia lutea, Johannsen, Ann. Ent. Soc. Amer. vol. 36, p. 781, 1943. 
Stilobezzia mallochinoSmSin, Ent. ^ews, vol. Z5, p. 2SS, 1924, (c?, 9; New York, 

Pennsylvania). — Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943 

(eastern United States). (New synonymy.) 

Description. — Female: Length 2.5 mm., wing 1.9 mm. by 0.7 mm. 
Yellow, including anterior wing veins, antennae, and legs ; mesonotum, 
except humeri, and apex of abdomen rufous brown. Vestiture of 
very long coarse brown bristles, those on scutellum as long as breadth 
of scutellum. 

Male: Ninth sternite nearly three times as broad as long, with a 
very broad, shallow, posterior emargination, the membrane spiculate; 
ninth tergite conical, with a prominent, submedian pair of long, fleshy, 
setose lobes. Basistyles simple, twice as long as broad; dististyles 
slightly longer than basistyles, curved, and gradually attenuated, 
with pointed apices. Aedeagus with a pair of oblique sinuate 
sclerites, bases of which are articulated with ventral root of basistyle, 
apices close together, pointed, and bent ventrad. Parameres with 
stout basal apodemes, the stems close together and slightly sinuate, 
flattened and slightly expanded past middle, with apices pointed and 
somewhat curved ventrad. 

Type. — In collection Illinois Natural History Survey, 9, Illinois, 
Elizabeth. 

Material examined. — New Hampshire: Center Harbor, July 9, 
Dyar, 1 9; Franconia, Slosson, 1 cT. 

232991—53 2 



66 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Connecticut: Green Falls, June 31, 1935, Chapman, 1 cf , 1 9. 

Rhode Island: Westerly, June 10, 1936, Chapman, 1 <^. 

New York: Armonk, Bemus Pomt, Canadarago Lake, Kast 
Bridge, Millwood, Poughkeepsie, Rome, June, July 1934-36, H. K. 
Townes, 12 cfd", 7 99. 

New Jersey: Moorestown, June 6, 1936, H. and M. Townes, 1 9; 
Riverside, June 18, 1939, H. K. Townes, 1 cf . 

Delaware: Smyrna, June 8, 1937, Bradley, 1 cf. 

Maryland: Cabin John, June 18, 1920, Aldrich, 1 9; Glen Echo, 
July 1, 1923, Malloch, 1 9; Plummer's Island, June 18, 1914, Shannon, 
1 9. 

District of Columbia: Rock Creek Park, June 17, 1920, Aldrich, 1 cf. 

Virginia; Dead Run, June 6, 1914, Shannon, 1 9; Falls Church, 
June 7, 1914, Shannon, 2 99, and July 4, 8, 1950, Wirth, 4 cT" cT, 7 99; 
Glencarlyn, June 7, 1935, Malloch, 1 cf , 1 9. 

South Carolina: Greenville, June 4, 1933, H. K. Townes, 1 cf. 

Indiana: Lafayette, June 26, 1916, Aldrich, 1 cf , 4 99. 

Illinois: Elizabeth, July 7, 1917, Malloch, 1 9 (type of Hartomyia 
lutea Malloch), 

Michigan: Cheboygan County, July 17, 1942, Sabrosky, 1 9. 

Iowa: Maquoka Caves State Park, Jackson Count}'', Jul}' 3, 1949, 
Laffoon, 1 9; Pikes Peak State Park, Clayton County, July 4, 1949, 
Laffoon, 1 9; Sioux City, June, July 1949-50, Slater and Laffoon, 
6 d'c?', 11 99. 

Remarks. — Alalloch had misidentified the male of this species as 
Hartomyia gilva (Coquillett), which is an Atrichopogon, in 1915, and 
in 1918 described the female of the same species as Hartomyia lutea, 
new species. Hoffman discovered that Malloch's determination of 
gilva was incorrect and described the species as Siilohezzia mallochi, 
1924, in part from Malloch's material. Since lioffman made no 
mention of Malloch's description of hitea, he ma}^ have been unaware 
of it, or he may not have been able to recognize Malloch's species. 
The present synonymy has been made after an examination of the 
type of lutea and specimens determined as gilva by IVfalloch in the 
Illinois Natural History Survey through the kindness of Dr. H. H. 
Ross, and of specimens in the U. S. National Museum determined by 
Hoffman as mallochi. Hoffman's detailed description of this species 
needs no elaboration except for the characters of the male genitalia. 

4. Stilobezzia {Neostilobezzia) stonei, new species 

Figure 11, c 

Diagnostic characters. — A large brownish species; mesonotum and 
scutellum dull yellowish ; legs yellowish; midfemora, hind femora, and 
tibiae dark brown; wings grayish brown; hal teres dull yellow. 



HELEID MIDGES, GENUS STILOBEZZIA — ^WIRTH 67 

Description. — Female: Length 1.8 mm., wing 1.8 mm. by 0.7 mm. 
Head dull brownish, including antennae and palpi, with a few long 
black hairs. Antenna as long as head and thorax combined; flagellar 
segments in proportion of 20:15:15:15:15:16:17:18:25:25:25:25:30. 
Palpal segments slender, in proportion of 5:10:12:6:10, third segment 
slightly swollen on basal two-thirds, with a small pit at distal fourth. 

Thorax dull brown, mesonotum and scutellum dull yellowish, 
postscutellum and pleura dark brown. Mesonotum with moderate 
to long brown hairs in rows; scutellum with about ten long brown 
marginal hairs. Legs dull brown, usually all of forelegs, midtarsus 
and hind tarsus yellowish. Proportions of segments of hind leg as in 
table 1; legs moderately slender, hairs fine, not very long except 
those on hind tibia about twice the diameter of segment; basi tarsi 
with ventral spines, three or four on basal half on forelegs and midlegs, 
one at base on hind leg; claws unequal, rather strong and curved, the 
longer nearly as long as fifth segment, the other about half as long. 

Wings evenly infuscated, the veins slightly darker, a few macro- 
trichiae at apices of cells Rg and Mi. First radial cell large, about a 
third the length of second; costa extending to 0.8 wing length; cross- 
vein r-m faint in middle, oblique; petiole of media as long as crossvein. 
Halteres yellow. Abdomen dark brown with short fine hairs; two 
large, slightly unequal ovoid spermathecae and a vestigial third 
present, ducts not sclerotized. 

Male: Similar to the female but with the usual sexual differences; 
antennal plumes brownish, knees quite dark. Ninth sternite about 
three times as broad as long, with broad posterior excavation more 
than halfway to base, the membrane bare; ninth tergite rounded, with 
small, submedian, setose, apicolateral lobes. Basistyles simple, twice 
as long as broad ; dististyles 0.9 times as long as basistyles, slender and 
nearly straight, their apices slightly knobbed. Aedeagus with an 
oblique pair of slightly curved sclerites with distal ends close together, 
sharp-pointed, and bent ventrad; apparently bearing a pair of hyaline 
membranous lobes on posterior margins and attached to membrane 
connecting dorsal roots of basistyles on ventral side of parameres. 
Parameres with broad, platelike, basal apodemes; stems very broad 
and flattened, gradually expanded distad with a short lateral point and 
a longer, slenderer, ventrally bent, distal point, their distal margins 
obliquely truncated. 

Types.— VSNM 60964, holotype, d", and aflotype, Falls Church, 
Va., July 8, 1950, Wirth. Paratypes: Virginia: 16 cTcf , 24 99, same 
data as type. Georgia: 2 cf c?", 17 99, Thomasville, May 1949, Palmer. 
Florida: 2 99, Innerarity Point, May 4, 1950, Eathert; 1 9, Panama 
City Beach, May 6, 1949, McElvey. 



68 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Remarks. — Related to S. lutea (Malloch) in general features, but 
readily distinguished by the more extensive brown coloration, 
finer vestiture and the characteristic broad, obliquely truncated, two- 
pointed, platelike, male parameres. This species is named in honor of 
Dr. Alan Stone. 

Subgenus Stilobezzia Kieffer, 1911 

Stilobezzia KiefiFer, Rec. Indian Mus., vol. 6, p. 118, 1911, and vol. 9, p. 184, 1913; 
Bull. Ent. Soc. France, p. 192, 1919. — Goetghebuer, Mem. Mus. Hist. Nat. 
Belgique, vol. 8, p. 59, 1920. — Carter, Ingram, and Macfie, Ann. Trop. Med. 
Parasit., vol. 15, p. 324, 1921. — Edwards, Trans. Ent. Soc. London, vol. 74, 
p. 411, 1926; Notulae Ent., vol. 9, p. 9, 1929.— Johannsen, Arch. Hydrobiol., 
SuppL, vol. 9, p. 430, 1931; Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943.— 
Ingram and Macfie, The Diptera of Patagonia and southern Chile . . ., 
pt. 2, fasc. 4, p. 191, 1931. — de Meillon, Proc. Ent. Soc. London, ser. B, vol. 
7, p. 266. — Tokunaga, Philippine Journ. Sci., vol. 72, p. 155, 1940. — Lane, 
Rev. Ent., vol. 18, p. 197, 1947.— Lee, Proc. Linn. Soc. New South Wales, 
vol. 72, p. 345, 1948.— Wirth, Univ. California Publ. Ent., vol. 9, p. 202, 1952. 
(Genotype, Ceratopogon notatus de Meijere, as Stilobezzia f estiva KiefiFer; 
original designation.) 

Hartomxjia Malloch, Bull. Illinois State Lab. Nat, Hist., vol. 10, p. 339, 1915. 
(Genotype, Ceratopogon pictus Coquillett; original designation.) 

Diagnosis. — Two radial cells present; wing without macrotrichiae 
at apex. 

5. Stilobezzia (Stilobezzia) antennalis (Coquillett), 1901 

Figure 12, e 

Ceratopogon antennalis Coquillett, Proc. U. S. Nat. Mus., vol. 23, p. 606, 1901 

( 9 , District of Columbia). 
Ceratolophiis antennalis, Kieffer, Genera insectorum, fasc. 42, p. 60, 1906. 
Johannseniella antennalis, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, 

p. 227, 1914. 
Hartomyia antennalis, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 343, 

1915 (cf ; Illinois, Indiana). 
Stilobezzia antennalis, Johannsen, Ann. Ent. Soc. Amer., 36: 781, 1943. 

Description. — Female: Length 1.5 mm., wing 1.6 mm. by 0.6 mm. 
Head, thorax, and abdomen polished black; antennae and legs yellow, 
midfemur and hind femur black except at extreme bases. Wings bare 
and hyaline, anterior veins yellowish, knob of hal teres black. Palpi 
and last five flagellar segments slightly infuscated, flagellar segments 
in proportion of 18:11:11:11:11:11:11:15:25:25:25:25:30. Palpal 
segments in proportion of 4:8:14:8:15, third segment not swollen, 
pit absent. Proportions of segments of hind leg as in table 1 ; basitarsi 
without spines, fifth segment with a pair of long ventral batonnets; 
claws long, simple, and very unequal, the short claw fused with base 
of and about a third as long as the other. First radial cell about a 
fourth as long as second; r-m crossvein vertical, petiole of media to a 



HELEID MIDGES, GENUS STILOBEZZIA — WIRTH 69 

third the length of Mi. Spermathecae two, the larger oval, the 
smaller subspherical and a third the diameter of other, a minute 
vestigial third present. 

Male: Similar to the female but with the usual sexual differences; 
plumes of antennae brown. Ninth sternite a narrow transverse ante- 
rior band, the membrane spiculate; ninth tergite evenly rounded 
caudad, the apicolateral lobes not prominent. Basis tyle with large 
irregular lobe on inner side to distal third; dististyle rather strongly 
bent past middle and tapered to pointed tip. Aedeagus with a pair 
of oblique, straight, sclerotized bars with distal apices scarcely meet- 
ing. Parameres a pair of strongly sclerotized, large, strongly sinuate 
rods with pointed apices and large basal knobs. 

Type.—USNM 5481, 9, District of Columbia. 

Material examined. — New York: Ithaca, June 11, 1935, H. K. 
Townes, 1 9; Rome, June 24, 1934, H. K. Townes, 1 d'. 

New Jersey: Moorestown, June 6, 1939, H. and M. Townes, 1 c?. 

Maryland: Dorchester, July 10, 1907, Barber, 1 9; Mayo, July 22, 
1950, Wirth, 2 cf cf, 1 9; Plummer's Island, June 30, 1914, Shannon, 

1 9, and Aug. 18, 1913, Viereck, 1 9. 

District of Columbia: June, 1 9 (type) ; Benning's, July 21, Aldrich, 

2 99. 

Virginia: June IG, 1903, Pergande, 2 cfcf, 3 99; Dead Run, June 
22, 1915, Shannon, 1 9, Falls Church, July 4, 8, 1950, Wu'th, 3 c^ d', 
4 99. 

Michigan: Midland County, July 17, 1944, Dreisbach, 1 9; Traverse 
City, June 17, 1943, Sabrosky, 1 cf. 

lUinois: Freeport, July 4, 1917, Malloch, 3 cf cf; Galena Junction, 
July 8, 1917, Malloch, 1 9; White Heath, June 4, 1939, Dirks, 1 9. 

Tennessee: Ivnoxville, May 25, Aldrich, 1 9. 

Mississippi: West Point, Aug. 11, 1904, Barber, 1 cf. 

Louisiana: Baton Rouge, May 4, 16, 1947, Wirth, 4 cf cf, 8 99. 

6. Stilobezzia {StUobezsia') beckae, new species 

Figure 11, / 

Diagnostic characters. — A large, dark brown species with silvery 
pruinose pattern on mesonotum and abdomen; wing with two darker 
and one lighter spots on anterior margin; knobs of hal teres black; 
mesonotum with strong anterior tubercle; fifth tarsal segment 
unspined. 

Description. — Female: Length 2.5 mm., wing 1.8 mm. b}^ 0.7 mm. 
Head black with dense whitish pruinosity, antennae brown, bases of 
flagellar segments 3 to 10 yellowish; palpi dark brown. Flagellar 
segments of antenna in proportion of 16:12:15:20:20:20:20:22:28: 



70 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

30:32:32:42. Palpal segments in proportion of 5:12:15:8:12, 
third segment slightly swollen, with large, shallow sensory pit near 
apex. 

Thorax dark reddish brown, with dense silvery pruinescence in 
irregular patches; mesonotum long, with prominent, conical, anterior 
spine, the anterior portion markedly sloping, posterior portion flat- 
tened between wing bases; covered with sparse, long, brown hairs; 
scutellum with four long, fine, brown hairs. Legs dark brown, all 
femora and fore tibiae with obscure broad median light bands, mid- 
tibia and hind tibia with narrow sub-basal light rings, tarsi yellow 
except fifth segment of each brown. Proportions of segments of 
hind leg as in table 1 ; legs with moderately long, black hairs, wliich are 
stronger on hind tibia ; basitarsi with three ventral spines on foreleg, 
four on midleg, and one at base on hind leg; fifth segment without 
ventral batonnets; claws very slender and slightly unequal, the longer 
as long as fifth segment, the other about 0.6 to 0.8 as long. 

Wing hyaline, without macrotrichiae, veins infuscated, a large 
brownish spot over first radial cell from vein Ri to crossvein r-m; 
apex of second radial cell with bro^vn spot, then a short white area 
with apex of wing weakly infuscated. First radial cell well developed, 
one-fourth as long as second ; r-m crossvein faint in middle, forming an 
oblique line with Ri, petiole of media as long as crossvein. Knobs of 
halteres black. 

Abdomen black, distal half of tergites 1 to 3 and narrow distal 
margins of remaining tergites with silvery, pruinose bands. Sper- 
mathecae two, subequal, pyriform. 

Male: Similar to the female but with the usual sexual differences; 
antennal plumes golden. Ninth sternite over twice as broad as long, 
with caudal excavation to half its length, the membrane bare; ninth 
tergite rounded with a pair of membranous, setigerous, apicolateral 
lobes. Basistyles simple, about 2.5 times as long as broad; dististyles 
slender, nearly as long as basistyles, gently curved, with pointed 
incurved apices. Aedeagus with a pair of heavj^, oblique, crooked, 
lateral sclerites meeting mesad subapically, with apices pointed and 
abruptly bent ventrolaterad. Parameres with heavy, platelike, 
V-shaped, basal apodemes; stems straight and contiguous mesad, 
slightly expanded midway, with lateral wall folded ventrad and mesad 
and apparently fusing on midline on distal third; extreme apices 
slender, flattened dorsoventrally and abruptly bent ventrad. 

Types. — USNM 60695, holotype, cf, and allotype. Crystal River, 
Citrus County, Fla., June 7, 1950, Hudson, light trap. Parat3T)es: 
Florida: 10 cf cf , 17 99, same data as type except dates, June 5-July 17, 
1949-50; 6 99, Grayton Beach, Walton County, May 10, 1949, Butler; 
15 99, Santa Rosa, Walton County, June 7, 1949, Peterson. Missis- 
sippi: 4 99, Horn Island, June, July, 1944, Richmond. 



HELEID MIDGES, GENUS STILOBEZZIA — ^WIRTH 71 

Additional material examined. — Peru: Iquitos, March, April 1931, 
Shannon, 1 9. 

Remarks. — Closely allied to S. paulistensis Lane, 1947, from Brazil 
which differs, however, in having the legs yellowish rather than black- 
ened, in lacking the dark spot at apex of second radial cell, and in 
having the lateral sclerites of the male aedeagus shorter, with apices 
capitate and bent mesad rather than slender and bent laterad. S. 
rabelloi Lane, 1947, is also closely related, but is a larger, hairier 
species with three dark wing spots, including one at apex of first 
radial cell, and has several pairs of batonnets on fifth tarsal segment. 
I am happy to name this species for Mrs. Elisabeth C. Beck of Arling- 
ton, Fla., who has kindly sent me so many interesting Florida heleids. 

7. Stilobezzia {Stilobezzia) hicolor Lane, 1947 

Figure 11, i 
Stilobezzia hicolor Lane, Rev. Ent., vol. 18, p. 208, 1947 (c^, Brazil). 

Description. — Female: (Here described for the first time.) Length 
1.5 mm., wing 1.5 mm. b}^ 0.6 mm. Head dark brown, antennal 
pedicel, clypeus, and palpi yellowish. Thorax and abdomen polished 
black, with metallic bluish violet reflections. Mesonotum rather 
broad and flat, with sparse, long, brown hairs, scutellum with four 
marginal bristles. Alidcoxa and hind coxa black, rest of legs yellow, 
except knees and fifth tarsal segments dark. Legs with fine hairs; 
proportions of segments as in table 1 ; basitarsi unspined ; fifth segment 
with a pair of long black batonnets at base: claws slender and unequal, 
the outer claw as long as fifth segment, the inner about a third as long. 
Wing hyaline, with violet reflections, anterior veins yellow; macro tri- 
chiae absent. First radial cell distinct, a sixth as long as second; 
petiole of media about twice as long as crossvein r-m. Halteres black. 
Abdomen convex above, somewhat petiolate, with a row of long, brown 
hairs across each tergite. 

Male: As in the female but with the usual sexual differences; 
plumes of antennae brown. Ninth sternite a narrow anterior band, 
the posterior membrane spiculate; ninth tergite rounded, with incon- 
spicuous, rounded, setigerous, apicolateral lobes. Basistyle with a 
small lobe at half the length of inner margin; distist3de about as long 
as basistyle, tapered to slender tip. Aedeagus with an only slightly 
oblique pair of sclerotized bars, stout in the Texas specimen figured, 
but slenderer in the tropical specimens. Parameres with lateral 
apodemes bent caudad; stems straight, stout and rodlike, with beak- 
like apices abruptly bent laterad. 

Type. — In coUection University of Sao Paulo, Brazil, cf , Brazil, 
Sao Paulo, Osasco. 



72 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 




g. ELEGANTULA 



h. COQUILLETTI 



I. BICOLOR 



Figure 11. — Male genitalia of Stilobezzia species: a, S. (S.) rabelloi Lane; b, S. (N.) lutea 
(Malloch); c. S. {N.) stonei, new species; d, S. (S.) punctipes, new species; e, S. {N.) 
fuscula Wirth;/, S. (S.) beckae, new species; g, S. (£.) elegantula (Johannsen); h, S. (S.) 
coquilletti Kieffer; i, S. (S.) bicolor Lane. 



HELEID MIDGES, GENUS STILOBEZZIAWIRTH 



73 




a. THOMSENAE 




b. BULLA 





c. SYBLEAE 




d, VIRIOIS 



e. ANTENNALIS 




/. DIVERSA 



g. PRUINOSA 



h. GLAUCA 



Figure 12. — Male genitalia of Stilobezzia species: a, S. (S.) thomsenae, new species; b, S. 
(S.) bulla Thomsen; c, S. (S.) sybleae, new species; d, S. (S.) viridis (Coquillett); e, S. (S.) 
antennalis (Coquillett); /, S. (S.) diversa (Coquillett); g, S. (S.) pruinosa Wirth; h, S. 
(S.) glauca Macfie. 



74 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Material examined. — Texas: San Antonio, Apr. 8, 1907. Pratt, 1 cf . 
Costa Rica: Higuito, Sdiild, 2 d' d', 10 99. 

Puerto Rico: Anasco, Apr. 22, 1933, Faxon, Mills, and Anderson, 
1 cf , 4 99. Bayamon, Jan. 28, 1934, Lesesner and Anderson, 1 9. 
Panamd: Summit, Canal Zone, November 1946, Krauss, 2 cT cf , 1 9. 

8. Stilohezzia {Stilobezzia) bulla Thomsen, 1935 

Figure 12, & 

Stilohezzia bulla Thomsen, Journ. New York Ent. Soc, vol. 43, p. 289, 1935 
(cf, 9 ; New York); Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943. 

Description. — Female. Length 1.0 mm., wing 0.8 mm. by 0.4 mm. 
Pruinose grayish brown; scutellum and legs yellowish ;hal teres white; 
wings clear, veins stramineous. Antennal segments in proportion of 
12:8:9:10:10:10:10:10:13:15:15:15:20. Palpi short and slender, 
segments in proportion of 5:7:7:5:7, third segment not swollen, pit 
absent. Basi tarsi and fifth segment without spines; claws simple, 
slender and unequal, the longer as long as fifth segment, the other 
a third as long. Wings clear, macrotrichiae absent. Spermathecae 
two, oval, slightly unequal, with ducts sclerotized a very short 
distance. 

Male: Ninth sternite a narrow anterior ribbon, the membrane 
spiculate; ninth tergite tapered to a pair of prominent, slender, setose, 
apicolateral lobes, truncate between then* bases. Basistyle slender, 
with a prominent, heavily sclerotized, platelike lobe articulating with 
the median sclerite (? of aedeagus); dististyle about half as long as 
basistyle, rather stout and nearly straight, tip bluntly pointed, with 
a distinct subapical tooth on ventral face of outer edge. Aedeagus 
with basal sclerites reduced to very small, narrow bars, connected at 
right angles by a pair of short, barlike sclerites to a more dorsal, large, 
heavily sclerotized, transverse sclerite shaped like a pair of buffalo 
horns, with points articulating with inner lobes of basistyles. Para- 
meres with slender, hook-shaped, basal apodemes, stems rodlike, 
slender and nearly straight, gradually tapered to long, simple, fine 
tips reaching nearly to apices of lobes of ninth tergite. 

Types. — In collection Cornell University, holotype, cf , and allo- 
type. New York, Ithaca, McLean Bogs. 

Material examined. — Maryland: Dorchester, July 10, 1907, Barber, 
1 d^. 

District of Columbia: Bennings, July 21, Aldrich, 1 9. 

Virginia: Falls Church, July 4, 8, Aug. 6, 1950, Wirth, 18 c^cf, 
24 99. 

Georgia: Thomasville, May 15-30, 1949, Palmer, 1 9. 

Florida: Crystal River, Sept. 18, 1950, Hudson, 1 9. 



HELEID MIDGES, GENUS STILOBEZZIA — WIRTH 75 

Louisiana: Baton Rouge, Apr. 19, 1947, Wirth, 1 cf ; Kilbourne, 
May 10, 1947, Wii-th, 1 9. 

9. Stilobezzia {Stilobezzia) coquilletti Kieffer, 1905 

Figure 11, h 

Ceratopogon pidus Coquillett, Journ. New York Ent. Soc, vol. 13, p. 60, 1905 

(9, Virginia). 
Hartomyia picta, Malloch, 1915, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 

341, 1915 (c?, Illinois). 
Stilobezzia coquilletti Kieffer, Ann. Mas. Nat. Hungarici, vol. 15, p. 308, 1917 

(new name for C. picta Coquillett not Meigen). — Johannsen, Ann. Ent. Soc. 

Amer., vol. 36, p. 781, 1943. 

Description. — Female: Length 2.0 mm., wing 2.0 mm. by 0.8 mm. 
Yellow, in some specimens with greenish tinge; with a pair of widely 
separated, narrow brown lines from anterior median point of meso- 
notum to ends of scutellum; also, narrow center of scutellum, palpi, 
mesopleiu"on and midcoxa, narrow apices of hind femur and tibia, 
fifth tarsal segment, and fore, lateral, and hind margins of abdominal 
tergites 3 to 7, brown. Wing bare and hyaline, all veins brownish, a 
brown spot over first radial cell including Ri and r-m crossvein; a 
second spot over apex of second radial cell and below halfway across 
cell R5; knobs of halteres brown. 

Flagellar segments of antenna in proportion of 25 :15: 15:15:15:15: 
17:18:28:28:30:30:60. Palpal segments in proportion of 4:15:20: 
10:22, third segment not swollen, pit absent. Mesonotum and 
abdomen with coarse, dark bristles. Proportions of segments of hind 
leg as in table 1 ; basitarsi without spines, fifth segment with a pair of 
long, black batonnets at basal third; claws slender and unequal, the 
longer as long as fifth segment, the other about half as long. Sper- 
matheca one, large and ovoid, the duct not sclerotized. Wing with 
macro trichiae absent; first radial cell very small and narrow, about a 
seventh as long as second, latter to 0.8 wing length; crossvein r-m 
forming an oblique line with vein Ri ; media petiolate for 0.23 its length. 

Male: Similar to the female but with the usual sexual differences; 
body more or less greenish with black markings as in female , antennal 
plumes yellow at bases, black on distal halves. Ninth sternite a 
narrow, transverse, anterior band, posterior membrane spiculate; 
ninth tergite full and rounded, with a pair of low, rounded, setigerous, 
apicolateral lobes. Basistyle about twice as long as broad at base, ab- 
ruptly narrowed about midway on inner side, forming a low, mesal lobe 
connected by an irregular membrane to ventrolateral margins of ninth 
tergite; dististyle about 0.7 as long as basistyle, stout, slightly incurved, 
with bluntly pointed tip. Aedeagus with an oblique pair of spatulate 
sclerites with bases abruptly bent laterad around bases of basistyles. 
Parameres a pair of broad, flattened, sclerotized plates, bases connected 
across mid^^ne by a highly sclerotized band from outer anterolateral 



76 PROCEEDINGS OP THE NATIONAL MUSEUM vol. 103 

margins of basistyles; base of each paramere with a prominent lateral 
spur projecting caudad along inner margin of basistyle; distal half 
greatly expanded, with a large, sharp, lateral tooth; inner margins 
contiguous at apices, each with a very slender, pointed spine abruptly 
bent ventrad and cephalad. 

Type.— VSNM 8356, 9, Virginia. 

Material examined. — Maryland: Shadyside, July 6, 1925, Aldrich, 
1 9. 

Virginia: June 16, Pergande, 1 9 (type). 

Illinois: Urbana, May 20, 1914, Aldrich, 1 cf, 19. 

Louisiana: Baton Rouge, Apr. 13, 25, 1947, Wirth, 3 cf cf , 3 99. 

Mexico: Ciudad Monte, Tamaulipas, Nov. 23-Dec. ], 1943, 
Brookman, 29 cT cf, 35 99; Tamazanchale, San Luis Potosi, March 11, 
1944, Brookman, 1 9; Tapachula, Chiapas, Sept. 20, 1944, Brookman, 
1 9. 

Puerto Rico: Arecibo, Apr. 5, 1932, Faxon and Anderson, 1 cf , 
1 9; San Juan, June 14, 1933, Harlan, 1 9. 

Jamaica: Bath, St. Thomas, Chapin and Blackwelder, 4 99; Spanish 
Town, Feb. 2, 1937, Chapin and Blackwelder, 3 99. 

Panama : Sabanas, Apr. 20, 1923, Shannon, 1 9. 

Remarks. — S. fiehrigi Kieffer, 1917, from Paraguay and Brazil, 
resembles S. coquilletti very closely except for the abdominal markings, 
with a pair of large lateral brown spots on the first tergite and a large 
median brown spot on the second, and tergites 3 to 7 with dark markings 
on the disc rather than on the margins. In the male genitalia of 
jiehrigi the basistyles bear a prominent mesal lobe, the dististyles are 
slenderer at the apices, the lateral bars of the aedeagus are slender 
without capitate tips, and the parameres are not so broad and the 
lateral tooth is not so prominent. 

10. Stilobezzia (Stilobezzia) diversa (Coquillett), 1901 

Figure 12, / 

Ceratopogon diversus Coquillett, Proc. U. S. Nat. Mus., vol. 23, p. 607, 1907 ( 9 , 

New Jersey). 
Ceratolophus diversus, Kieffer, Genera insectoruui, fasc. 42, p. 60, 1906. 
Johannseniella diversa, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 

227, 1914. 
Hariomyia diversa, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 344, 

1915. 
Stilobezzia diversa, Johannsen, Ann. Ent. Soc. Anier., vol. 36, p. 781, 1943. 

Description. — Female: Length 1.5 mm., wing 1.4 mm. by 0.6 mm. 
Head brown, antennae and palpi yellow; thorax shining black; abdo- 
men and coxae light green; legs yellow, fifth tarsal segment brownish; 
wings hyaline, anterior veins yellow, halteres j^ellowish. Antennae as 
long as entire body; palpi slender, third segment not swollen. Wing 



HELEID MIDGES, GENUS STILOBEZZIA — WIRTH 77 

with first radial cell a fifth as long as second; petiole of media a fourth 
as long as distal portion. Legs slender, nearly bare, proportions of 
segments of hind leg as in table 1 ; basitarsi unspined, fifth segment 
with a pair of ventral batonnets. 

Male: Similar to the female but with the usual sexual differences; 
plumes of antennae yellow. Ninth sternite about four times as broad 
as long, with a very shallow, broad, posterior excavation, the posterior 
membrane spiculate; ninth tergite tapered, much longer than basi- 
styles, with very short, setigerous, apicolateral lobes. Basistyles broad 
at bases, attenuated distad, simple, with an irregular, hyaline mem- 
brane connecting the inner margins about two-thirds way to apices; 
dististyles about as long as basistyles, slightly curved and tapered to a 
slender, pointed tip. Aedeagus with a pair of slender, sclerotized, 
oblique rods with straight apices, the curved bases continuous with 
anterior margins of basistyles. Parameres with bases very stout and 
conical, and with greatly narrowed apices recurved ventrolaterad, the 
bases connected by a narrow, sclerotized band. 

Type.— Vii'NM 5482, 9, New Jersey, Riverton. 

Material examined. — New Jersey: Riverton, July 3, Johnson, 1 9 
(type). 

Virginia: Falls Church, July 4, Aug. 6, 1950, Wu'th, 4 cf cf , 3 99. 

Georgia: Thomasville, May 15-30, 1949, Palmer, 1 cf , 2 99. 

11. Stilobezzia (Stilobezzia) glauca Macfie, 1939 

Figure 12, h 

Stilobezzia glauca Macfie, Rev. Ent., vol. 10, p. 204, 1939 (cf, Brazil). — Lane, Rev. 
Ent., vol. 18, p. 207, 1947 ( 9 ). 

Diagnostic characters. — A rather small, bright, pale green species 
with whitish antennae, legs, wings, and halteres; mesonotum exten- 
sively dark, shining brown; abdomen with black band across third 
tergite and lateral spots on fifth. 

Description. — Female: Length 1.8 mm., wing 1.7 mm. by 0.6 mm. 
Head yellowish brown, antennae whitish at bases, last five segments 
and palpi pale brownish. Flagellar segments in proportion of 
25:15:15:15:16:17:18:20:40:40:35:35:45. Palpi long, segments in pro- 
portion of 4:7:15:8:12; third segment not swollen, pit absent. 

Thorax pale green; mesonotum with median third of anterior margin 
shining brownish black, forming a broad, V-shaped mark caudad, the 
sides of mesonotum broadly dark to ends of scutellum, and the median 
area between the dark bands yellowish. Vestiture consisting of a few 
long, brown bristles; scutellum with four weak, marginal bristles. 
Postscutellum and a faint pleural spot brown. Legs whitish, hind 
femur and tibia with very faint dark, apical spot. Proportions of 
segments of hind leg as in table 1 ; basitarsus without spines, fifth seg- 



78 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

ment with a pair of long, dark batonnets at basal fourth; claws very 
slender and unequal, the longer claw as long as fifth segment, the other 
a third as long. 

Wings hyaline, anterior veins yellowish; first radial cell small, a 
seventh as long as second; petiole of media a fifth way to apex of wing. 
Halteres yelloMash white. 

Abdomen pale green; two dark spots present on sides of first and 
second tergites; third tergite with broad, black band; fifth tergite 
with a pair of small, lateral, black spots. Spermatheca one, slightly 
oval, the duct not sclerotized. 

Male: Similar to the female but with the usual sexual differences ; 
antennal plumes bright yellow. Ninth sternite broad and not strongly 
sclerotized, with spiculate membrane; ninth tergite rounded with 
membranous, setigerous, apicolateral lobes. Basistyle with distal 
portion slender, base stout, a large unsclerotized lobe about halfway 
along inner margin, connected on dorsal side to margin of ninth tergite ; 
dististyle about as long as basistyle, tapered and curved to a slender, 
bluntly pointed tip. Aedeagus with an oblique pair of long, straight, 
slender, sclerotized bars. Parameres with stout basal knobs, a pair 
of inner apodemes joined mesad by a membrane from their anterior 
margins; stems stout, slightly curved, with apex abruptly narrowed, 
pointed, and bent mesad in the form of a short claw. 

Type. — In collection British Museum (Natural History), cf, 
Brazil, Nova Teutonia. 

Material examined. — Virginia: Falls Church, July 4-29, 1950, 
Wu-th, 9 cf d', 16 99. 

South Carolina: Greenville, June 20, 1940, H. and M. Townes, 1 cf . 

Louisiana: Baton Rouge, May 1947, Wirth, 2 cfcf. 

Remarks. — Very similar to S. viridis (Coquillett), 1901, in the gen- 
eral green color with black-banded abdomen, but readily distinguished 
from that species by the dark-banded mesonotum, unhanded legs, 
pale wings, and in the male by the yellowish antennal plumes. One 
female from Falls Church is atypical, with the abdomen entirely dark. 

12. Stilobezzia {Stiloheszia) pallidiventris (Mallocb), 1915 

Hartomyia pallidiventris Malioch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 

344, 1915 (9 ; Illinois, Indiana). 
Stilobezzia pallidiventris, Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943. 

Description. — Female (from original description): Length 1.0 mm. 
Shining black; head, antennae, palpi, legs, and abdomen yellow; 
abdomen may be darkened; wings hyaline; halteres white. Antennae 
as long as entire body; legs slender; basitarsi longer than segments 
2 to 5 combined; fifth segment without ventral batonnets; claws 
slender, inner half as long as outer. First radial cell a fifth as long as 
second. 



HELEID MIDGES, GENUS STILOBEZZIA — WIRTH 79 

Type. — In collection Illinois Natural History Survey, 9, Illinois, 
Urbana. 

Remarks. — Very near S. (S.) diver sa (Coquillett), but much smaller, 
the abdomen yellowish rather than green, and fifth tarsal segment of 
female without ventral batonnets. 

13. Stilobeszia (Stilobezzia) pruinosa Wirth, 1952 

Figure 12, g 

Stilobezzia pruinosa Wirth, Univ. California Publ. Ent., vol. 9, p. 203, 1952 (cf, 
9 ; California). 

Description. — Female: Length 1.0 mm. Pruinose brown, with 
broad, short, unmarked wings and a marked, pruinose gray pattern 
on the mesonotum; legs with narrow sub-basal pale rings on tibiae; 
halteres white. Proportions of segments of hind leg as in table 1; 
basitarsi and fifth segment without ventral spines. Spermathecae 
two, slightly unequal, slightly ovoid, with a few minute perforations; 
the ducts sclerotized a very short distance. 

Male: Ninth sternite transverse; ninth tergite long and rounded 
caudad with a subapical pair of low, rounded, setigerous lobes. Basi- 
styles tapered, the inner margins with low hump bearing a few setose 
tubercles; dististyles very stout, with blunt apices. Aedeagus with 
a pair of small, oblique, sclerotized bars. Parameres a pair of highly 
sclerotized, heavy plates; basal apodemes winglike and bilobed; 
inner margins of stems nearly straight, slightly expanded on outer 
sides about halfway to apices; apices abruptly narrowed and bent 
ventromesad on about distal fourth. The apices of the parameres 
are somewhat longer in the Florida specimens figured than in the type 
from California. 

Types. — USNM 59945, holotype, cf, allotype, California, San Luis 
Obispo County, Arroyo Seco Ranger Station. 

Material examined. — California: AiToyo Seco Ranger Station, 
San Luis Obispo County, July 1, 1948, Wirth, 3 cf cf (type, 2 para- 
types), 1 9 (allotype); Pollack Pines, Eldorado County, July 7, 1948, 
Wirth, 1 cf ; Snowline Camp, Eldorado County, July 8, 1948, Wirth, 
1 (f . 

Florida: Gainesville, Alachua County, June 2, 1950, Bidlingmayer 
1 cf , 1 9; Grayton Beach, Walton County, May 10, 1949, Butler, 1 9 

14. Stilobezzia {Stilobezzia") punctipes, new species 

Figure 11, d 

Diagonostic characters. — A large, yellowish species with brown 
thorax and brown-punctate legs; abdomen with narrow segmental 
dark bands; wings with two small black spots. 



80 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i03 

Description. — Female: Length 2.4 mm., wing 1.9 mm. by 0.9 mm. 
Head pruinose brown; antennae yellow, apices of segments 3 to 10 
and aU of last five segments brown; palpi black. Flagellar segments 
in proportion of 28:18:18:18:18:18:20:20:38:38:40:40:55, last 
segment with very slender pointed tip. Papal segments in proportion 
of 5:10:16:8:20, third segment slightly swollen with a small, round, 
subapical pit. 

Thorax pruinose brown, margins of mesonotum, broad center of 
scatellum, and upper half of pleiu-a yellow. Mesonotum with irregular, 
silv^ery pruinose patches, a few long, brown bristles; scutellum with 
four long, marginal bristles. Coxae and trochanters dark brown, 
rest of legs yellow; fore femur with broad sub-basal, midfemur with 
narrow basal, and hind femur with narrow apical brown bands; all 
tibiae narrowly brown at apices; rows of small, round, brown puncta- 
tions at bases of hairs on all femora and tibiae, these hairs long and 
dark on hind tibia. Proportions of segments of hind leg as in table 
1; basitarsus with three black ventral spines on forelegs, two on 
midlegs and one at base on hind legs; fifth tarsal segment without 
batonnets; claws slender and unequal, the longer claw as long as 
fifth segment, the other two-thirds as long. 

Wing hyaline, anterior veins yellowish; a brown spot over r-m 
crossvein and a very small one over vein R2+3; macrotrichiae absent. 
First radial cell large, one-fourth as long as second, latter to 0.8 wing 
length; petiole of media as long as r-m crossvein. Halteres brown. 

Abdomen dull, pruinose yellowish, tergites with irregular brown 
bands in middle of segments, pleura dark. Spermathecae three, the 
two larger subequal and oval with short, sclerotized ducts, the third 
about two-thirds as large and subspherical, without sclerotized duct. 

Male: Similar to the female but with the usual sexual differences; 
plumes of antennae golden brown. Ninth sternite a very narrow band, 
the posterior membrane spiculate; ninth tergite rounded, with small, 
submedian, setigerous lobes at apex. Basistyle simple, twice as long 
as broad; dististyle as long as basistyle and rather stout to apex, 
slightly bent on distal third. Aedeagus with an oblique pair of heavy 
twice-sinuate sclerites with apices snoutlike and crossing each other 
mesad. Parameres very slender, the lateral apodemes evenly curved 
and slender, stems nearly straight with bases slightly enlarged, 
gradually tapered to sharp, hooked tips abruptly bent ventrocephalad 
at two-thirds total length. 

Ti/pes.— USNM 60966, holotype, 9, Crystal River, Citrus County, 
Fla., June 7, 1949, Hudson, light trap; allotype, same data except 
date, Oct. 6, 1949. Paratypes: Florida: 11 d'cf, 23 99, same data 
as type except dates, May-October 1949-50; 1 cT, 3 99, Leesburg, Lake 
County, Aug. 7, 1949, Braddock; Mexico: 4 99, Ciudad Monte, 
Tamaulipas, Nov. 22, 1943, Brookman. 



HELEID MIDGES, GENUS STILOBEZZIA — ^WIRTH 81 

Most closely related to the Neotropical species S. (S.) punctulata 
Lane, 1947, and S. (S.) kiefferi Lane, 1947, which it resembles in its 
spotted wings and punctate legs. Both related species, however, 
have the abdomen reddish, the former species differing in having 
three dark wing spots, the third over tip of second radial cell, while 
the second species has the first radial cell half as long as the second, 
yellowish coxae, whitish halteres, and the fore femur and midfemur 
lack the basal dark bands. 

15. Stilobezzia {Stilobezzia) rabelloi Lane, 1947 

Figure 11, a 

Stilobezzia rabelloi Lane, Rev. Ent., vol. 18, p. 203, 1947 (c?, 9 ; Brazil). 

Diagnostic characters. — A large, brown species with irregular, 
silvery, pruinose markings; wing brownish, infuscated with tlu-ee 
darker and one lighter spots on anterior margin; body hairs dense and 
strong; fifth tarsal segment with 3 or 4 pairs of ventral batonnets. 

Description. — Female. Length 3.0 mm., wing 2.2 mm. by 0.8 mm. 
Head pruinose brown, narrow bases of flagellar segments yellow, 
palpi black. Antenna with flagellar segments in proportion of 
22:15:15:15:18:20:24:24:40:42:50:?:? (last two segments broken, 
not measured). Palpal segments in proportion of 8:15:25:15:20, 
third segment slightly swollen, densely set with strong hairs, a small, 
round pit near apex. 

Mesonotum with strong, conical, anterior spine, the sides sloping, 
flattened only in prescutellar area; color dark brown with irregular 
silvery pruinose markings, usually three sublateral pairs of small, 
rounded, polished spots on disc; humeral corners yellowish; a few long, 
brown hairs in row. Scutellum brown, yellowish in middle, with 
4 or 5 long, brown, marginal bristles. Postscutellum and pleura dark, 
pollinose brown. Legs including coxae dark brown; fore femur and 
midfemur with narrow sub-basal and subapical and broad median 
yellowish bands; hind femur with narrow, light band at apex; fore 
tibia and midtibia with narrow sub-basal and broad median yellowish 
bands; first four tarsal segments yellowish. Proportions of segments 
of hind leg as in table 1 ; leg hairs strong and spinose, especially long 
on hind tibia; basitarsus with strong black basal and apical spines, a 
third spine in middle on forelegs; fifth segment with 3 or 4 pairs of 
ventral batonnets; claws very slender and unequal, the longer claw 
as long as fifth segment, the other a third as long. 

Wing brownish hyaline, extensively infuscated along the veins; 
three dark brown spots, first over r-m crossvein, second over vein 
R2+3, and the third over distal third of second radial cell and extending 
halfway through cell Rj at same level; a light area just beyond tip 



82 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

of costa as large as third browTi spot, the apex of the wing beyond 
Hghtly infuscated. First radial cell nearly half as long as second, the 
latter ending at about 0.7 wing length; petiole of media about as long 
as first radial cell. Halteres brown except at extreme bases of stems. 

Abdomen dark brown, tergites with narrow, apical, segmental, 
silvery pruinose bands; vestitiu-e very dense, consisting of long, stout, 
brown hairs. Spermathecae two, ovoid, subequal, with ducts sclero- 
tized a short distance, and a rudimentary third present. 

Male: Similar to the female but with the usual sexual differences; 
plumes of antennae bro^\^l. Ninth sternite about three times as 
broad as long, with shallow, mesal, posterior excavation, and the 
membrane bare; ninth tergite short and rounded with very short, 
setigerous, apicolateral lobes. Basistyles simple, very long, slender 
and tapered to tips, about three times as long as broad; dististyles 
about 0.8 as long as basistjdes, slightly incurved, slender, with pointed 
apices. Aedeagus with an oblique pair of double-sinuate sclerites 
with mesal tips snoutlike. Parameres strongl}^ sclerotized with 
broad, winglike, lateral apodemes at bases, a lateral pair of L-shaped 
or boomerang-shaped lateral plates with blunt apices directed latero- 
caudad, and a submedian pair of nearly straight processes of same 
thickness, but about twice as long, with pointed apices. 

Type. — -In collection University of Sao Paulo, Brazil, cf, Brazil, 
Rio de Janeiro, Estrada Rio-Sao Paulo, km. 47. 

Material examined. — Georgia: Thomasville, ]May 15-30, 1949, 
Palmer, 1 9. 

Florida: Leesburg, Lake County, Aug, 7, 1949, Braddock, 2 99; 
Panama City Beach, May 6, 1949, McElvey, 3 cf cf , 1 9; Santa Rosa 
Island, Escambia County, June 7, 1949, Butler, 1 cf , 1 9. 

Louisiana: New Orleans, May 27, 1947, Van Beeck, 6 cf cf . 

Honduras: La Ceiba, Aug. 6, 1916, Dyer, 1 cf . 

Surinam: Moengo, Nov. 10, 1947, Stage, 1 9. 

Remarks. — Closely related to S. (S.) beckae, new species, which it 
resembles closely in the bro\vn mesonotum with silvery pruinosity and 
strong anterior spinose tubercle and brownish infuscated and spotted 
wings, but beckae differs in lacking the third dark spot at apex of first 
radial cell and the ventral batonnets on fifth tarsal segment, and the 
body hairs are not nearly so well developed. 

16. Stilobezzia {Stilobezzia) sybleae, new species 

Figure 12, c 

Diagnostic characters. — A small, shining, brown species with dusky, 
yellow legs; wing hyaline with a prominent beadlike swelling of vein 
^i+5 near apex of second radial cell; hind tarsi with fifth segment and 
claws greatly reduced. 



HELEID MIDGES, GENUS STILOBEZZIA — WIRTH 83 

Description. — Female: Length 1.5 mm., wing 1.3 mm. by 0.5 mm. 
Head shining brown, clypeus and antenna! pedicel yellowish, antennal 
flagellum and palpi dusky. Thorax polished brown, mesonotum with 
sparse black hah's in rows ; scutellum with four marginal hairs. Pleura, 
coxae, midfemur, hind femur, and tibiae dusky, yellowish brown; all 
of forelegs, midtarsus, and hind tarsus lighter yellowish. Proportions 
of segments of hind leg as in table 1 ; basi tarsi unspined; fifth segment 
with a pair of black ventral batonnets near base; claws slender and 
unequal, normal on forelegs and midlegs with outer claw as long as 
fifth segment, the inner a third as long, on hind legs with fifth segment 
half as long and claws only a fourth as long as on forelegs and midlegs. 

Wings grayish hyaline, without macro trichiae, anterior veins brown- 
ish yellow. First radial cell a fourth as long as second, latter to five- 
eighths wing length ; vein R4+5 on distal half of second radial ceU with 
an oval beadlike swelling; cross vein r-m short, petiole of media about 
three times as long as crossvein. Halteres brown. Abdomen shining 
brown, very convex dorsally and somewhat petiolate at base. 

Male: Ninth sternite a slender anterior band, the posterior mem- 
brane spiculate; ninth tergite rounded, the apicolateral lobes reduced. 
Basistyle broad, with a large triangular, flattened, sclerotized lobe 
about midway of mesal margin; dististyle nearly as long as basistyle, 
incurved, gradually tapered to slender apex. Aedeagus with a pair of 
slightly oblique, long, very slender, sclerotized bars. Parameres with 
basal knoblike apodemes fused with anteromesal margins of inner 
lobes of basistyles, with slender, long, curved, sicklelike posterior 
blades. 

Types.— USNM 60967, holotype, 9, and allotype, Falls Church, 
Virginia, July 4, 1950, Wirth. 

Remarks. — This species is very similar to *S'. (S.) antennalis (Coquil- 
lett), but the highly modified posterior branch of the radius and hind 
fifth tarsal segment and claws are quite distinctive. The West 
African species S. limnophila Ingram and Macfie, 1922, is very similar 
in coloration and in the structure of the male genitalia, but since the 
female is unloiown, the wing and tarsal features cannot be compared 
with those of sybleae. Parabezzia poikiloptera Ingram and Macfie, 
1922, from West Africa, which is a Stilobezzia of the subgenus Eukrai- 
ohelea, has a remarkable thickening of the radius almost exactly like 
that of sybleae, indicating that this character is not just an aberration. 
This species is named for my wife, Syble Austin Wirth. 

17. Stilobezzia {Stilobezzia) thomsenae, new species 

Figure 12, a 

Diagnostic characters. — Very closely related to, and almost indis- 
tinguishable from, bulla Thomsen, except for the very characteristic 
male genitaha. 



84 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Description. — Female: Length 1.0 mm.; wing 0.8 mm. by 0.4 mm. 
Apparently somewhat darker brown than bulla, the mesonotum and 
scutellum concolorous; femora brownish. Otherwise as in bulla. 

Male: Genitalia broader than long, parts very broad and stout. 
Ninth sternite a narrow, anterior band, posterior membrane spiculate; 
ninth tergite rounded, with a submedian pair of fingerlike setose lobes 
about twice as long as broad. Basistyles simple, about twice as long as 
broad, set far apart, no inner lobes; dististyles short and stout, two- 
thirds as long as basistyle, with abruptly narrowed, pointed, clawlike 
apices. Aedeagus with anterior arms vestigial, the dorsal median 
sclerite a crescent-shaped, transverse bar connecting distoventral 
angles of basistyles. Parameres stout, with broad, lateral, winglike, 
basal apodemes, stems shghtly knobbed at bases, stout and slightly 
out-curved toward apices, which are slightly flattened, expanded, 
rounded, and bent ventrad. 

Type.—VSNM 609G8, holotype, cf, Everglades City, Collier 
County, Fla., Feb. 7, 1950, Davidson, light trap. 

Remarks. — One female from Crystal River, Citrus County, Fla., 
Sept. 18, 1950, Hudson, is provisionally referred to S. (S.) thomsenae, 
since it resembles the male in coloration, although it may prove to be 
a dark specimen of S. (S.) bulla. This species is named in honor of 
Dr. Lillian Thomsen, who has contributed much to the study of 
North American Heleidae. 

18. Stilobezzia {Stilobezzia) viridis (Coquillett), 1901 

Figure 12, d 
Ceratopogon viridis Coquillett, Proc. U. S. Nat. Mus., vol. 23, p. 607, 1901 ( 9 , 

New Jersey). 
Ceratolophus viridis, Kieffer, Genera insectorum, fasc. 42, p. 61, 1906. 
Johannseniella viridis, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 

227, 1914. 
Hartomyia viridis, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 342, 

1915. 
Stilobezzia viridis, Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943. 

Description. — Female: Length 2.0 mm., wing 1.9 mm. by 0.6 mm. 
A bright, pale-green species; antennae and legs yellow, apex of antenna 
dark; distal fourth of hind femur and narrow apex of hind tibia black; 
abdomen with a black band across distal half of third tergite and a 
pair of large black spots on fifth tergite. Wings brownish hyaline, 
anterior veins infuscated. Proportions of segments of hind leg as in 
table 1 ; basitarsi without spines ; fifth segment with a pair of long, 
black batonnets at basal fom-th; claw^s slender and very unequal, the 
longer as long as fifth segment, the other about half as long. 

Male: As in the female but with the usual sexual differences, 



HELEID MIDGES, GENUS STILOBEZZIA — ^WIRTH 85 

plumes of antennae dark. Genitalia very large and bulbous. Ninth 
sternite reduced, the posterior membrane spiculate; ninth tergite 
large, convex, apex rounded with low, rounded, setigerous, apicolateral 
lobes. Basistyle short and very broad, nearly as broad as long; 
dististyle about 1.2 times as long as basistyle, extreme base bulbous, 
distal portion very slender, irregularly bent inward, with blunt apex 
slightly bent outward. Aedeagus with a pair of slender, nearly 
straight, barlike sclerites. Parameres with reduced basal scleroti- 
zation, stems slender and slightly crooked, apices greatly expanded in 
a rounded lobe. 

Type.—USNM 5483, 9, New Jersey, Riverton. 

Material examined. — New Jersey: Riverton, June 16, Johnson 
(type). 

Maryland: Cabin John, May 26, 1943, Cortes and Townes, 2 cf cf ; 
Glen Echo, June 30, 1929, Bridwell, 1 cf . 

Florida: Leesburg, Lake County, Aug. 7, 1949, Braddock, 1 9. 

Texas: Kirbyville, May 20, 1908, Tucker, Icf. 

Remarks. — S. glauca Macfie from Brazil and the southeastern 
United States is closely related to viridis, but has the antennae and 
wings infuscated, the mesonotum dark, and lacks the apical black 
markings on the hind femur. 

References 

GOETGHEBUER, M. 

1934. Heleidae (Ceratopogonidae). In Lindner, Die Fliegen der Palal- 
arktischen Region, Lief. 78, pp. 49-94. 

JOHANNSEN, 0. A. 

1934. New species of North American Ceratopogonidae and Chironomidae. 

Journ. New York Ent. Soc, vol. 42, pp. 343-353. 
1943a. Two new species of American Ceratopogonidae (Diptera). Ann. 

Ent. Soc. Amer., vol. 36, pp. 761-762. 
1943b. A generic synopsis of the Ceratopogonidae (Heleidae) of the Americas, 

a bibliography, and a list of the North American species. Ann. 

Ent. Soc. Amer., vol. 36, pp. 763-791. 

1946. Some new species of Nemocerous Diptera from Guam. Bull. Bishop 

Mus., No. 189, pp. 189-193. 
Lane, J. 

1947. Especies Brasileiras de Slilobezzia (Dipt., Ceratopogonidae) e Zygone- 

ura stonei nov. nom. (Dipt., Mycetophilidae) . Rev. Ent., vol. 18, 
pp. 197-214, 9 figs. 
Macfie, J. W. S. 

1935. Ceratopogonidae (Dipt.) from the river Amazon. Stylops, vol. 4, 

pp. 49-56, 2 figs. 
Meillon, B. de 

1938. Notes on African Ceratopogonidae (Diptera). Proc. Ent. Soc. 
London, ser. B, vol 7, pp. 266-270. 

U. S. eoVEKNMENT PRINTINS OFriCEi IMS 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 




SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 



Vol. 103 Washington: 1953 No. 3317 

BEETLES OF THE OEDEMERID GENUS VASACES 
CHAMPION 



By Ross H. Arnett, Jr.^ 



Champion (1889) described the genus Vasaces for three species from 
Central America. One species, Vasaces aeneipe'nnis , was described 
from material from Mexico, and the other two, Vasaces sordidus and 
Vasaces costatus, were described from Guatemala. 

Since the completion of my revision of the Nearctic Oedemeridae 
(Amer. Midi. Nat., vol. 45, pp. 257-391, 1951), I have seen several 
specimens of Vasaces collected by J. N. Knull in Arizona and Texas. 
This genus has not previously been recorded from North America, 
north of Mexico. In addition, a few specimens from Central America, 
in the collections of the U. S. National Museum and the Museum of 
Comparative Zoology, have been available to me for study. 

The present paper, based on all available material, with the excep- 
tion of the Champion type specimens, adds four new species to the 
genus. A key to the species is included, and the position of the genus 
in relation to other oedemerid genera is discussed. 

I wish to thank Mr. J. N. Knull, Ohio State University, and Dr. 
P. J. Darlington, Museum of Comparative Zoology, for the loan of 
material used in this study. 

'Bureau of Entomology and Plant Quarantine, U. S. Department of Agriculture. 

232714—53 87 



88 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Genus Fasaces Champion, 1889 

Vasaces Champion, Biologia Central!- Americana, Coleoptera, vol. 4, pt. 2., p. Ill, 
1889 [no included species] and p. 128, pi. 6, figs. 7 and 7, a-c, 1890 [first in- 
cluded species]. — Arnett, Journ. Washington Acad. Sci., vol. 40, p. 225, 1950. 
(Genotype, Vasaces aeneipennis Champion, 1890; designated by Arnett, 
1950.) 

This genus belongs in the subfamily Oedemerinae as treated in my 
paper on the Nearctic species. The apical segment of the maxillary 
palpus is triangular or cultriform and the fore tibia has two apical 
spurs. It is tentatively placed in the tribe Asclerini, as I have pre- 
viously defined it. However, studies of certain other genera have 
led me to revise somewhat my views on the oedemerid tribes, and 
when more groups have been restudied some rearrangement may be 
necessary. Vasaces Champion will key to couplet 7 of my key to the 
genera of Asclerini. The slightly emarginate eyes separate it from 
Heliocis Arnett, and the triangular or cultriform apical segment of the 
maxillary palpus will separate it from the genus Sisenes Champion. 
In Sisenes Champion the apical segment of the maxiUary palpus is 
small, hardly triangular. In addition, many Sisenes species resemble 
lycids and have the antennal segments somewhat flattened, a condi- 
tion which is not found in Vasaces. In the key to the male genitalia, 
this genus runs to couplet 3, where it may be separated from the 
genera Eumecomera Arnett and Sisenes on the basis of the structure 
of the ninth abdominal sternite. In Vasaces the apical portion of the 
ninth sternite of both Eumecomera and Sisenes consists of two separate 
rods united only by thin membrane. The ninth sternite of Vasaces 
resembles somewhat that of Heliocis except that the base is two 
distinct rods instead of the Y-shaped piece found in Heliocis. 

It appears that the genus Sisenes is a close relative of Vasaces, 
both on characters of external morphology and those of the male 
genitalia. However, in the studied forms, the two genera can be 
adequately separated using the characters mentioned above. In 
gross appearance, the two genera are very distinct. The species of 
Vasaces are for the most part dark, somewhat metallic, elongate 
beetles, quite different from the brilliantly colored, lycidlike species 
of Sisenes. 

Generic diagnosis. — Head moderate or elongate, usually constricted 
behind the eyes. Eyes somewhat emarginate, large. Antenna 
slender, 11-segmented, inserted between the eye and the base of the 
mandible, distinctly separated from the margin of the eye and the 
base of the mandible. Second segment of the antenna large, over 
one-half the length of the fourth and sometimes nearly the length of 



OEDEMERID BEETLES — ^ARNETT 89 

the third. Both mandibles with the apex of each bifid. Apical 
segment of the maxillary palpus large, triangular or cultriform, widest 
at the base or near the center. Thorax slightly longer than broad, 
usually only slightly expanded apically and only slightly constricted 
at the base. Legs slender, claws simple; all tarsi with only the 
penultimate segment of each spongy beneath. Elytra distinctly 
costate, sometimes markedly so. 

Male genitalia: Median lobe short, base wider, without proc- 
esses, tapering towards apex, without apical enlargements. Paramere 
a troughlike plate, wide basally, without basal processes, narrowed 
toward apex; apex V-shaped, emarginate; tegminite large, spatulate. 
Ninth abdominal sternite hood-shaped, with two basal tails; apex 
expanded, with basally projecting lateral flanges. Ninth tergite 
two parallel rods, apically expanded and united by membrane. 
Eighth abdominal sternite and tergite each bilobed, the sternite larger. 
Setae present on the apical lobes of the eighth abdominal segment. 

Biology. — Unknown. 

Distribution. — KJno^vn only from Central America and the extreme 
southwestern part of the United States. 

Key to the species of Vasaces 

1. Apical segment of the maxillary palpus cultriform (fig. 13, gr), its outer edge 

sinuate 2 

Apical segment of the maxillary palpus triangular (fig. 13, /i) 4 

2. Costae, suture, and margins of elytra fusco-piceous, intervals pale; antennae 

uniformly dark except for small pale areas at the joints; legs dark, 

slightly paler at joints linearis, new species 

Elytra with uniform coloration 3 

3. Elytra uniformly brassy costatus Champion 

Elytra dark knulli, new species 

4. Pale testaceous; elytra with costae and suture each with an indistinct, inter- 

rupted row of fine, oblong streaks on either side sordidus Champion 

Dark species, otherwise marked 5 

5. Pronotum orange with black maculations, head short__maculatus, new species 
Pronotum dark, head elongate 6 

6. Pronotum with sides nearly parallel elongatus, new species 

Pronotum distinctly sinuate laterally aenipennis, Champion 

Vasaces linearis, new species 
FiGUBB 13, c, e-g, j~l, m, n 

Diagnostic characters. — The light brassy color, with the dark brown, 
prominent elytral costae, readily differentiates this species from the 
others. It is most closely related to V. costatus since the shape of 
the apical segment of the maxillary palpus is similar. 



90 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i03 

Description. — (Based on male holotype.) Antenna long, nearly as 
long as the body, second segment subequal to the third. Apical 
segment of the maxillary palpus large, distinctly cultriform, outer 
margin arcuate. Vertex of head transversely elevated between the 
eyes, and with a slight median groove from the base of the antenna 
to the summit of the vertex. Pronotum somewhat longer than broad, 
widest near anterior margin, slightly narrower at base, surface irregu- 
lar, laterally with slight elevations, median line somewhat impressed, 
and with a slight V-shaped elevation on anterior half. Elytron long 
and narrow, with four prominent costae and the suture and margin 
prominently elevated, and a short, basal, spurious costa uniting with 
the inner margin. 

Entire insect clothed with fine, short, sparse, whitish hairs. Color: 
in general pale brassy; head with a darker area on front ; mouthparts, 
except for maxillary palpi, paler. Antennae and maxillary palpi dark. 
Thorax dark except for irregular paler areas near base and apex. 
Elytra uniformly pale brassy except for dark brown costae, suture, 
and margins. Ventral surface and legs dark. Surface irregularly 
rugose-punctate. Length: 8 mm. 

Male genitalia: Described in the generic diagnosis. The seventh 
sternite has a slight angular emargination which is probably a second- 
ary sexual character. 

Type. — Holotype, male, Ohio State Univ. coll., Arizona, Huachuca 
Mountains, Cochise County, June 9, 1935, J. N. Knull. 

Distribution. — Known only from this single male type specimen 
from Arizona. 

Vasaces costatus Champion, 1890 

Vasaces costatus Champion, Biologia Centrali-Americana, Coleoptera, vol. 4, 
pt. 2, p. 129, pi. 6, fig. 8, 1890. 

This species is also known only from one male, the type, from 
Guatemala, at Cerro Zunil (4,000 feet), collected by Champion, and 
is in the British Museum (Natm-al History). It is much smaller 
than the genotype, but a little larger (9.5 mm.) than the preceding 
species. The elytra are uniformly brassy in color, lacking the dark 
brown costal, sutural, and marginal markings. The seventh sternite 
of the male is deeply emarginate. 

Vasaces knulli, new species 

Figure 13, b 

Diagnostic characters. — The small size, dark color, and lack of 
brassy tints serve to separate this species from the other two having the 
apical segment of the maxillary palpus cultriform. It is most closely 
allied to V. costatus. 



OEDEMERID BEETLES — ^ARNETT 



91 




Figure 13. — Adults and male genitalia of the genus V as aces: a, Vasaces maculatus, new- 
species, 9 ;b, F. knulli, new species, 9 ; c, F. linearis, new species, cf ; d, V. elongatus, new 
species, ? , ^, /, cf genitalia, V. linearis, new species: e, paramere;/, median lobe; g, h, 
maxillary palpi: g, V. linearis, new species, c^; h, V. elongatus, new species, 9 ; ^, V- 
aeneipennis Champion, 9 ; j-n, V. linearis, new species, cf ;y, ninth tergite; k, ninth ster- 
nite; /, tegminite; m, eighth tergite; n, eighth sternite. 



92 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

Description. — (Based on female holotype.) Antennae short, not 
extending over two-thirds the length of the elytra, second and third 
segments equal in length, each about three-fourths the length of the 
fourth segment. Apical segment of the maxillary palpus large, 
cultriform, outer edge slightly arcuate. Head shorter than that of the 
other species; front with a slight depression near the bases of the 
antennae on the inner side; a moderately deeply impressed median 
line on the front. Pronotum with surface very irregular and with a 
moderately prominent V-shaped ridge on the anterior half and 
an interrupted lateral ridge on each side; longer than broad, widest 
anteriorly just beyond the middle, constricted behind. Elytra nar- 
row, long; costae present but obscure. Surface covered with fine, 
short, sparse, whitish hairs. Color: dark, mouthparts, underside 
of head, neck, coxae, bases of femora, tarsi at joints, and small area at 
joints of the segments of the antennae paler. Surface finely rugose- 
punctate. Length: 7 mm. 

Type. — Holotype, female, Ohio State Univ. coll., Arizona, Huachuca 
Mountains, Cochise County, June 9, 1935, J. N. Knull. 

Distribution. — This species is known only from the type, which is 
from Arizona. 

Vasaces sordiduSf Champion, 1890 

Vasaces sordidus Champion, Biologia Centrali- Americana, Coleoptera, vol. 4, pt. 
2, p. 129, 1890. 

This species is known from one female only, the type, from CapetUlo, 
Guatemala, collected also by Champion, which is in the British Mu- 
seum (Natural History). It is smaller than the preceding species 
(8.5 mm.) ; the apical segment of the maxUlary palpus is smaller, and 
triangular instead of cultriform. The color is pale testaceous. The 
costae and suture are conspicuous, and each has an indistinct, inter- 
rupted row of fine oblong streaks on either side. 

Vasaces maculatus, new species 

FiGUBE 13, a 

Diagnostic characters. — The dark orange pronotum with a median and 
two lateral spots is unique among the species of this genus. It is the 
largest known species in the genus and is most closely related to V. 
sordidus but has the orange markings which are lacking in that species. 

Description. — (Based on female holotype.) Antennae short, reach- 
ing to about two-thirds the length of the elytra, second segment short, 
about two-thirds as long as the third. Apical segment of the maxUlary 
palpus large, elongate triangular, widest near basal third, outer mar- 
gin straight. Front of head irregular, with a circular depression 
extending from the inner side of the base of the antenna basad about 
half the distance to the pronotum. Pronotum quadrate, widest 



OEDEMERID BEETLES — ARNETT 93 

anteiiorly, only slightly constricted posteriorly, siu'face irregular but 
without prominent ridges or depressions except for a slight depression 
on either side near the anterior lateral angle. Elytra long, narrower 
apically, about twice as broad as thorax; costae present, flat, not promi- 
nent. 

Entire insect clothed with very fine, short, sparse, golden hairs. 
Surface finely rugose-punctate. Head with punctures larger, more 
distinct, with a stouter recumbent hair arising near each. Color: 
Head orange, with a large dark spot on front; antennae and palpi dark; 
thorax orange except for a large central dark spot which connects with 
two large lateral spots ; legs with coxae and bases of femora pale, re- 
mainder dark ; abdomen and elytra uniformly dark. Length : 1 mm . ; 
of paratype, 11 mm. 

Types. — Holotype, female, USNM [61112, Arizona, Chiricahua 
Mountains, June 8. Hubbard and Schwarz. Paratype, female, same 
data as type except date, June 11. 

The paratype agrees with the type except that the black markings 
of the thorax are less extensive. 

Distribution. — Known only from the two type specimens from 
Arizona. 

Vasaces elongatits, new species 

Figure 13, d, h 

Diagnostic characters. — The uniform dark color, elongate head and 
thorax, and parallel sides of the pronotum are characteristic of this 
species. It is very different in general habitus from the other members 
of this genus, but is probably most closely related to V. aeneipennis. 

Description. — Head elongate, over twice as long as wide at bases 
of antennae, hardly constricted behind the eyes. Antennae reaching 
beyond apical third of elytra, apical segment of the maxillary palpus 
triangular, widest at middle, outer edge straight. Front of head 
smooth, without elevations or depressions. Pronotum longer than 
broad (7 :4) , surface irregular but without distinct ridges or depressions, 
sides only slightly sinuate. Elytra broad, twice the width of the 
thorax, tapering behind. Legs long and slender, costae distinct, flat. 

Entire insect clothed with short, fine, white hairs, those of the head 
and thorax sparser, but longer and coarser than those of the rest of 
the body. Surface of head smooth and shiny with large distinct 
punctures which are sparser on the vertex; thorax coarsely rugose; 
remainder of body finely rugose. Color: dark fuscous; neck, under 
surface of head, mouthparts, and coxae paler. Length: 8 mm. 

Type. — Holotype, female, Ohio State Univ. coll., Texas, Chisos 
Mountains, Brewster County, June 9, 1939, D. J. and J. N. KnuU. 

Distribution. — KJaown only from the female type. 



94 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

Vasaces aeneipennis Champion 

Figure 13, i 

Vasaces aenipenms Champion, Biologia Centrali-Americana, Coleoptera, vol. 4, 
pt. 2, p. 128, 1890. 

This is apparently the least rare of the species in this genus and 
is one of the largest of the known species. The triangular apical 
segment of the maxillary palpus, the elongate head, and the uniform 
brassy color distinguish it from the other species. 

Champion described this species from two specimens (a male and 
a female) from Totosinapan, Mexico, in the Salle collection (location 
unknown). The U. S. National Museum collection contains two fe- 
males from Guatemala, and the Museum of Comparative Zoology has 
one female from Honduras. A detailed description of one female 
from Guatemala follo^vs: 

Description. — (Based on female, from Guatemala). Head elongate, 
about twice as long as wide at bases of antennae. Antennae long, 
reaching nearly to apical third of the elytra, second segment about 
two-thirds the length of the third. Apical segment of the maxillary 
palpus triangular, widest at middle, outer edge straight. Dorsal 
surface of head hregular, but without distinct ridges or depressions. 
Pronotum nearly quadrate, widest anteriorly, sides distinctly sinuate, 
surface irregular. Elytra long and narrow, tapering at apex; costae 
distinct, flat, both edges of costae outlined by a very fine, irregular, 
impressed line, somewhat darker in color, giving in gross appearance 
the impression that the costae are double. 

Surface clothed with extremely fine, very short, sparse, golden 
hairs which are somewhat larger on the thorax and dorsal surface of 
the head. Surface rugose, that of the thorax and head indistinctly 
rugose-punctate. Color: uniformly brassy, dorsal surface of the head 
and ventral surface of the thorax and abdomen darker. Length: 
15 mm. 

Variation : The surface of the head and thorax varies from rugose- 
punctate to shiny smooth with sparse punctures. The length varies 
from 10 to 16 mm. The color is darker in some specimens than in 
others. 

Distribution. — Known only from the types from Mexico and the 
above-mentioned specimens from Guatemala and Honduras (Rosario 
San Juancito). 

Material examined.— Female, USNM coll., Guatemala, Santa 
Maria, W. Schaus. 



U. S, GOVERNMENT PRINTING OFFICE: 1953 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



issued ira»?fvA. sJ^Ml ^y '^fi 




SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 



Vol.103 Washington : 1953 No. 3318 

SCARABAEID BEETLES OF THE GENUS BRADYCINETULUS 
AND CLOSELY RELATED GENERA IN THE UNITED STATES 



By O. L. Cartwright 



The group of beetles reviewed in this paper have long been known 
under the name Bolboceras. The species found in the United States 
were last revised by Charles Schaeffer, 1906 (Trans. Amer. Ent. Soc, 
vol. 32, pp. 249-254). At that time five species were placed in Horn's 
genus Bradycinetus, later incorrectly synonymized under Bolboceras 
(see synonymy under Bradycinetulus) , and three were assigned to 
Bolboceras. Of these three, Scarabaeus lazarus Fabricius had been 
placed in Eucanthus by Westwood, 1848 (Proc. Linn. Soc. London, 
vol. 1, p. 387), and Bradycinetus hornii Rivers and B. minor Linell 
were later removed to Kolbeus by Chapin (see synonymy under 
Bolbelasmus) . 

With the exception of the species Eucanthus and Bolbocerosoma, here 
considered correctly placed in recent usage, all of the United States 
species listed by Schaeffer have been reexamined in this study and 
reassigned to genera as required by the literature on the group. Also 
included in this paper are the one species described siiice Schaeffer's 
time, Bolboceras angulus Robinson; a Mexican species discovered in the 
United States; another which may eventually be found inside our 
borders; and four new United States species. Two new genera are 
proposed. A key, based in part on that by Boucomont, is presented 
for the genera of the Bolboceratini found in the United States. The 
genera Kolbeus, Bolbelasmus, Bradycinetulus, and the two new genera 
Bolbocerastes and Bolborhombus are discussed, synonymy listed, and a 
key to the species given. Pertinent references, the type locality and 
location of the type, and notes on distribution follow for each species. 
Distribution is also shown on the maps, figures 15 and 16. The genus 

232990—53 1 95 



96 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Bolbocerosoma, revised by Dawson and McColloch, 1924 (Canadian 
Ent., vol. 56, pp. 9-15), is not included, nor is the true Bolboceras, 
which was revised under the name Odontaeus by J. B. Wallis, 1928 
(Canadian Ent., vol. 60, pp. 119-128, 151-156, 168-176). 

The number of specimens available for study was greatly increased by 
material borrowed from museum and private collections. I am grate- 
ful for the opportunity to study these added specimens and thank the 
following who loaned museum specimens: R. H. Beamer, Snow 
Museum, University of Kansas (KAN); E, C. Becker, Canadian 
Department of Agriculture (CD A) ; M. A. Cazier, American Museum 
of Natural History (AMNH) ; P. J. Darlington, Museum of Compara- 
tive Zoology (MCZ) ; Henry Dietrich, Cornell University (CU) ; T. H. 
Hubbell, University of Michigan (MICH); P. D. Hurd, University 
of California (CAL) ; J. N. Knull, Ohio State University (OSU) ; Hugh 
B. Leech, California Academy of Sciences (CAS) ; L. M. Martin, Los 
Angeles County Museum (LAC) ; H. J. Reinhard, A. & M. College of 
Texas (TEX); V. D. Roth, Oregon State CoUege (ORE); M. W. 
Sanderson, Illinois Natural History Survey (INHS) ; and G. E. 
Wallace, Carnegie Museum, Pittsburgh (CARN). My thanks are 
extended also to those who loaned specimens from their private collec- 
tions: R. H. Arnett, D. K. Duncan, C. A. Frost, P. C. Grassman, 
Henry Howden, A. T. McClay, G. H. Nelson, F. H. Parker, Mark 
Robinson, E. C. VanDyke, and F. G. Werner. I am especially grate- 
ful to Henry Howden who examined my types, compared specimens 
with them and later furnished those records marked with an asterisk. 
The few records of the eastern Brady cinetulus ferrugineus (Palisot de 
Beauvois) taken from the literature are marked with a double asterisk. 
Florida records from the literature are from Blatchley (Florida Ent., 
vol. 12, p. 29, 1925), Alabama records from Loding (Geol. Surv. 
Alabama Monogr, 11, p. 101, 1945), and North Carolina records from 
Brimley (The insects of North Carolina, p. 201, 1938). 

AU other distribution records were copied from labels attached 
to specimens I examined. 

Key to the genera of Bolboceratini 

1 . Eye entirely divided by ocular canthus 2 

Eye not completely divided by ocular canthus 3 

2. Middle coxae narrowly separated; the intercoxal process with a vertical tooth- 

like elevation in front of the coxae; species usually bicolored. -Bolbocerosoma 

The intercoxal process without toothlike elevation; species not bicolored; 

males usually with long cephalic horn Bolboceras * 

3. Middle coxae subcontiguous ; metasternum linear between the coxae 4 

Middle coxae separated by the anterior lobe of the metasternal plate, which is 

never linear between the coxae 6 

> Odontaeus is a synonym. See Introduction and also discussion under Bradycinetulus. 



BOLBOCERATIlSn OF THE UNITED STATES — CARTWRIGHT 97 

4. Elytra with 5 discal striae Eucanthus 

Elytra with 7 discal striae Bolbelasmus 

5. Base of elytia margined, metasternal plate pyriform in outline, pronotum 

without postapical carina, first elytral stria interrupted by scutellum 6 

Base of elytra not margined, metasternal plate rhomboid in outline, pronotum 
with postapical carina, first two elytral striae interrupted by scutellum. 

Bolborhombus, new genus 

6. Apex of tibia of middle and hind legs deeply emarginate on outer side, the 

angle adjacent to spurs appearing almost as a fixed spur; without prosternal 

spine Bradycinetulus 

Apex of tibia of middle and hind legs obliquely truncate; prosternal spine 
behind anterior coxae transverse, doubly pointed, and remote from acutely 
angled intercoxal piece Bolbocerastes, new genus 

Genus Bolbelasmus Boucotnont, 1911 

Bolbelasmus Boucomont, Ann. Soc. Ent. France, vol. 79 (1910), p. 335, 1911. 
Kolbeus Boucomont, Ann. Soc. Ent. France, vol. 79 (1910), p. 335, 1911. — Chapin, 
Proc. Biol. Soc. Washington, vol. 59, p. 79, 1946. 

Chapin, 1946, placed Bradycinetus hornii Rivers and B. minor 
Linell in the genus Kolbeus, basing his action on their close relation- 
ship to Kolbeus arcuatus (Bates). Examination of specimens of these 
species in comparison with Bolbelasmus gallicus (Miilsant) and B. 
unicorne (Shrank), and in the light of Boucomont's key to the genera, 
places B. minor Linell and B. hornii Rivers in Bolbelasmus rather 
than in Kolbeus. 

The genus Kolbeus was erected without a genotype by Boucomont, 
1911, for the species Bolboceras arcuatus Bates and B. coreanus Kolbe. 
Lucas, 1920 (Catalogus . . . Coleopterorum, p. 356), selected 
B. coreanus Kolbe, a species found in Korea and China, as the 
genotype. The species B. arcuatus Bates almost exactly duplicates 
Bolbelasmus minor (Linell) except for the slightly narrower scutellum 
and lack of a marginal line over the middle half of the base of the 
pronotum. Since a direct comparison of specimens shows such minor 
differences, I transfer arcuatus Bates to the genus Bolbelasmus. 
Whether the genus Kolbeus should be synonymized with Bolbelasmus 
will depend upon an examination of its genotype, a species not avail- 
able for study at present. Further basis for placing arcuatus in Bol- 
belasmus is found in similarity of the aedeagi (see fig. 14). 

Genotype. — Scarabaeus gallicus Mulsant, Histoire naturelle des 
Coleopteres de France, Lamellicornes, p. 350, 1842. (Present desig- 
nation.) 

Key to the species of Bolbelasmus 

1. Base of pronotum not margined over middle half; length 7 to 9 mm. (Cali- 
fornia, Mexico) arcuatus (Bates) 

Base of pronotum completely margined ,-.,. 2 



98 PROCEEDINGS OF THE NATIONAL MUSEUM vol.103 

2. Ely tral striae coarsely, deeply punctate as in Eucanthus; genae obtusely roundfed ; 
surface of head and clypeus similarly densely, coarsely punctate in male; 
frontal carina weakly tri-tuberculate in female; length 7 to 10 mm. (Texas). 

minor (Linell) 

Ely tral striae finely to moderately punctate; genae angulate externally; 

surface of clypeus roughly, densely, coarsely punctate, remaining surface of 

head much smoother and more finely punctate in the male; frontal carina 

not trinodose in female; length 14 mm. (California) hornii (Rivers) 

Bolbelasmus arcuatus (Bates), new combination 

Figure 14, I 

Bolboceras arcuatus Bates, Biologia Centrali-Amerlcana, Coleoptera, vol. 2, pt. 2, 

p. Ill, 1887. 
Kolbeus arcuatus Boucomont, Ann. Soc. Ent. France, vol. 79 (1910), p. 336, 1911. — 

Chapin, Proc. Biol. Soc. Washington, vol. 59, p. 79, 1946. 

Bolbelasmus arcuatus (Bates) is a small, dark red-brown, shining 
species, 7 to 9 mm. in length, readily recognized through the lack of 
basal margins on both pronotum and elytra. The labrum is wddely, 
shallowly emarginate in front; the clypeus is almost evenly arcuate 
from side to side; the clypeal suture is distmct; the anterior angles of 
the pronotum are sharply right angled; the scutellum is elongate; 
the strial punctures are coarse, in the outside row at the apex very coarse 
and unusually deep, practically perforating the elytra; the elytra 
have a row of widely spaced, long, bristlelike hairs near the margin; 
the male has a single median conical cephalic horn and a low, wide, 
median thoracic prominence and anterior shallow depression on each 
side; the female has a wide, more-or-less tri-tuberculate carina on the 
head and a simple, fine, sharp, transverse carina on the pronotum. 

Type. — Type probably among the Biologia specimens in the British 
Museum (Natural History). 

Type locality. — Not designated among the localities originally listed, 
"Mexico, Cordova, Toxpan, Playa Vicente (Salle), Colima City 
(Hoge); Nicaragua, Chontales (Belt, Janson)." 

Specimens examined. — 19, 

Distribution. — Costa Rica: Tilaran (Guanacaste), San Pedro de 
Montes de Oca, San Jose, and *Dominical. Guatemala. Mexico: 
Colima, El Sabino, Compostela, MazatMn, Monterey, *Monclova, 
and *Guadalajara. United States: a single specimen labeled "Los 
Angeles, California, H. Klages Coll.," in the Carnegie Museum, 
Pittsburgh, Pa. 

Figure 14. — Ventral and lateral views of the aedeagi of Bradycinetulus and closely related 
species: a-c, Species of Bolborhombus: a, parvulus, new species; b, angulus (Robinson); 
c, schaefferi (Boucomont). d-g, Species of Bolbocerastes: d, peninsularis (Schaeffer); e, 
regalis, new species;! f, serratus (LeConte); g, imperialis, new species, h-j. Species of 
Bradycinetulus: h, ferrugineus (Palisot de Beauvois); i, rex, new species; j, fossatus 
(Haldeman). k-n, Species of Bolbelasmus: k, gallicus (Mulsant); /, arcuatus (Bates); 
m, hornii (Rivers); n, minor (Linell). 



BOLBOCERATINI OF THE UNITED STATES — CARTWRIGHT 99 




L ^ V y M fg 

Figure 14. — (For explanation see facing page.) 



100 PROCEEDINGS OF THE NATIONAL AIUSEUM vol. los 

Bolbelasmus minor (Linell), new combination 

Figure 14, n 

Bradycinetus minor Linell, Proc. U. S. Nat. Mus., vol. 18, p. 723, 1895. 
Bolboceras minor Schaeffer, Trans. Amer. Ent. Soc, vol. 32, p. 253, 1906. 
Kolbeus minor Chapin, Proc. Biol. Soc. Washington, vol. 59, p. 79, 1946. 

Bolbelasmus minor (Linell) almost exactly duplicates B. arcuatus 
(Bates) except that it averages slightly larger, the punctures are 
coarser throughout, and the pronotum is completely and distinctly 
margined basally. It is a small, dark red-bro-wn, shming species, 
7 to 10 mm. in length; the labrum is widely, shallowly emarginate in 
front; the clypeus is almost evenly arcuate from side to side; the 
clypeal suture is distinct; the anterior angles of the pronotum are 
sharply right angled; the scutellum is elongate; the strial punctures 
are very coarse, those at the apex of the elytra being tubular and 
so deep as to nearly perforate the elytra ; near the margin of the elytra 
is a row of widely spaced, long, bristlelike hairs; the male has the 
head and clypeus similarly densely punctate, the head bears a median, 
conical horn, and the pronotum has a low, wide, median prominence 
with anterior, shallow depressions each side; the female has a wide, 
transverse, tri-tuberculate carina on the head and a simple, fine, 
transverse caruia on the pronotum. 

Type.— Rolotyipe female, USNM 562. 

Type locality. — San Diego, Tex. 

Specimens examined. — 2 1 . 

Distribution. — Texas: Hidalgo County, Weslaco, Brownsville, Kings- 
ville, and San Diego. 

Season. — May 16 to June 16. 

BolbelasTnus hornii (Rivers), new combination 

Figure 14, m 

Bradycinetus hornii Rivers, Bull. California Acad. Sci., vol. 2, p. 61, 1886. 
Bolboceras hornii Schaeffer, Trans. Amer. Ent. Soc, vol. 32, p. 253, 1906. 
Bradycineiulus hornii Cockerell, Ent. News, vol. 17, p. 242, 1906. 
Kolbeus hornii Chapin, Proc. Biol. Soc. Washington, vol. 59, p. 79, 1946. 

Bolbelasmus hornii (Rivers) is quite similar to B. minor (Linell) but 
is considerably larger, lighter in color, and has much finer strial 
punctures. The scutellum is not so elongate, being nearly an equi- 
lateral triangle with arcuate sides. Other characters given in the 
key should readily identify this species. 

Type. — Location of type unknown to me. (According to Dr. 
VanDyke the entire Rivers Collection was purchased by Dr. Walther 
Horn and it is presumably in the Berlin-Dahlem collection.) 

Type locality. — "Sonora, Tuolumne Co., Cal. Found also in 
Sacramento Co." 



BOLBOCERATINI OF THE UNITED STATES — CARTWRIGHT 101 

Specimens examined. — 349. 

Distribution. — California: Sequoia National Park, Napa, Kaweah, 
Camp Greely (Tulare County), Mendocino, Mokelumne Hill, Camp 
Greely (Fresno County), Three Eiders, El Dorado County, Coulter- 
v^ille, Sonoma County, Potwisha, San Joaquin Experiment Station 
(Madera County), Mount Diablo, Sacramento, Mineralking, La Jon 
Pass, Kattlesnake (Placer County), *Cala7eras County, and *0'Neals 
(Madera County). Baja California: Hamilton Ranch. 

Season. — January 19 to May 8, with greatest number in April. 

Genus Brady cinetulus Cockerell, 1906 

Bolboceras Kirby (in part), Trans. Linn. Soc. London, vol. 12, pt. 2, p. 459, 1818. 

(Type not designated.) 
Amechanus Horn (not Thomson, 1864), Trans. Amer. Ent. Soc, vol. 3, p. 48, 

1870. 
Bradycinetus Horn (not Sars, 1865), Trans. Amer. Ent. Soc, vol. 3, p. 334, 1871. 
Bradycellus Schaeffer (not Erichson, 1837), Trans. Amer. Ent. Soc, vol. 32, p. 

249, 1906. 
Bradycinetulus Cockerell, Ent. News, vol. 17, p. 242, 1906. (Type not designated.) 

The genus Bolboceras was erected by Kirby in 1818 for the eight 
species Scarabaeus mobilicornis Linnaeus, S. mobilicornis vax. tes- 
taceus Fabricius, S. guadridens Fabricius, S. farctus Fabricius, S. 
lazarus Fabricius, S. cyclops Olivier, *S'. cephus Fabricius, and Bol- 
boceras australasiae Kirby. Technically no genotype was designated 
although he wrote, "My details of Bolboceras were taken from B. 
quadridens." Nearly all of the species mentioned have since been 
moved to other genera. Curtis, 1829 (British Ent., vol. 1, pt. 1, p. 
74), selected the species Scarabaeus mobilicornis Fabricius as the type 
of Bolboceras. Therefore, since our species of Odontaeus are con- 
generic with mobilicornis, they now take the generic name Bolboceras 
and it becomes necessary to find an available name for those species 
we have formerly placed in Bolboceras. A search of the literature 
indicates that name must be Bradycinetulus, as shown in the 
synonymy. 

Characters for Bradycinetulus, in addition to those listed in the key 
to genera, include the following: The mandibles are evenly arcuate ex- 
ternally; the median prominence of the pronotum is developed into a 
pair of horns in the male; the scutellum is wider than long and dis- 
tinctly punctate with fine to moderate punctures; and the posterior 
vertical face of the prosternal intercoxal piece is wide and flat, with 
the ventral edge evenly arcuate or slightly angulate at middle. The 
three species placed here are all large, 17 to 21 mm. in length by 10 
to 12 mm. in width. They are sufficiently differentiated in the follow- 
ing key. 

Genotype . — Scarabaeus ferrugineus Palisot de Beauvois, 1 809 . (Pres- 
ent designation.) 



102 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Key to the species of Bradycinetulus 

1. Eye canthus nearly straight edged laterally, wide clypeal horn of male bisinuate 

apically, the external angles projecting forward, acute, and scarcely thicker 
than the adjacent edges, which converge posteriorly; median thoracic horns 
heavy, directed upward, their sharply rounded tips recurved posteriorly 
(North Carolina to Florida and Mississippi). 

ferrugiiieus (Palisot de Beauvois) 
Eye canthus notched or emarginate, not straight edged laterally 2 

2. Wide clypeal horn of male distinctly trinodose apically, the external angles 

not projecting bej^ond median point, lateral edges parallel; median thoracic 
horns heavy, projecting forward and outward, sharply rounded and very 

little recurved (Nebraska to Texas and Arkansas) fossaius (Haldeman) 

Apex of clypeal horn of male widely truncate, straight across, the external 
angles slightly nodose, lateral edges parallel over apical two-fifths, then 
diverging to base; median thoracic horns slender, directed forward and up- 
ward, basally flattened in front and behind (Texas) rex, new species. 

Bradycinetulus ferrugineus (Palisot de Beauvois), 1809 

Figure 14, h: Plate 3 

Scarabaeus ferrugineus Palisot de Beauvois, Insectes recueillis en Afrique et en 

Amerique, livr. 6, p. 90, 1809. 
Bolboceras lecontei Dejean, Catalogue des Coleoptfeies . . ., ed. 3, vol. 3, p. 149, 

1833. 
Athyreus ferrugineus Klug, Abh. Berlin Acad., 1843, p. 22. 
Bolboceras ferrugineus Lacordaire, Histoire naturelle des insectes, vol. 3, p. 143, 

1856. 
Amechanus ferrugineus Horn, Trans. Amer. Ent. Soc, vol. 3, p. 48, 1870. 
Bradycinetus ferrugineus Horn, Ent. Amer., vol. 1, p. 89, 1885. 
Athyreus (Bradicinetus) ferrugineus Boucomont, in Wytsmanu, Genera insectorum, 

fasc. 7, p. 8, 1902. 
Bradycinetus ferrugineus Schaeffer, Trans. Amer. Ent. Soc, vol. 32, p. 250, 1906. 
Bradycinetulus ferrugineus Cockerell, Ent. News, vol. 17, p. 242, 1906. 
Bolboceras (Amechamus) ferrugineus Boucomont, Ann. Soc. Ent. France, vol. 79, 

p. 341, 1910. 

Type. — Location of type unknown to me. 

Type locality. — South CaroHna. 

Specimens examined. — 107. 

Distribution. — North Carolina: West End, Southern Pines, **Beau- 
fort, *Hamlet, *Carolina Beach, and **Tarboro. South Carolina: 
Meredith, Florence, Windsor, Johns Island, and Bulls Island. Geor- 
gia: Bainbridge, Spring Creek (Decatur County), and Augusta. Flor- 
ida: Port Saint Joe, Kissimmi, Orlando, Sanford, Enterprise, Miami, 
Crescent City, Indian River, Cedar Keys, *Lutz, *Wacissa, *Gaines- 
ville, **Saint Augustine, **LaGrange, and **Dunedin. Alabama: 
**Lee County, and **Mobile County. Mississippi: Ocean Springs, 
Biloxi, and Leakesville. 

Season. — March 11 to September 23, with the largest number of 
records in June and July. 



BOLBOCERATINI OF THE UNITED STATES — CARTWRIGHT 103 

Bradycinetulus fossatus (Haldeman), 1853 

Figure 14, j; Plate 3 

Bolboceras fossatus Haldeman, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 362, 

1853. 
Amechanus fossatus Horn, Trans. Amer. Ent. Soc, vol. 3, p. 48, 1870. 
Bradycinetus fossatus Schaeffer, Trans. Amer. Ent. Soc, vol. 32, p. 250, 1906. 
Bradycinetulus fossator Cockerell, Ent. News, vol. 17, p. 242, 1906. 
Bolboceras {Amechamus) fossatus Boucomont, Ann. Soc. Ent. France, vol. 79, 

p. 341, 1910. 

In a few male specimens of Bradycinetulus fossatus (Haldeman) the 
outer cariniform edges bordering the thoracic fossae show a slight 
tendencA^ to break near the upper end to form a second inner carina, a 
step tow^ard the condition found in Bolbocerastes. 

Type. — In LeConte collection, MCZ. 

Type locality. — Texas. 

Specimens examined. — 102. 

Distribution. — Nebraska: Imperial. Kansas: Reno County, Me- 
dora, and Sylvia. Oldahoma: Noble County, Cleveland County, 
Payne County, Thomas, Cleo Springs, Ferris, and *Alva. Arkansas: 
Pine Bluff. Texas: Fedor, Henrietta, Tyler, Jacksonville, Gold- 
thwaite, Robinson, Lexington, Jefferson, College Station, Lindale, 
Bexar County, Dallas, Victoria, Columbus, Paris, Austin, San Antonio, 
and Morris County. 

Season. — May 16 to August, with most records in June. 

Bradycinetulus rex, new species 

Figure 14, i; Plate 3 

Holotype male, length 21 mm., width 13 mm. In general appearance 
verj^ similar to Bradycinetulus jerrugineus (Palisot de Beauvois) and 
B. fossatus (Haldeman), rufo-testaceous, moderateh^ shining, densely 
hairy beneath. Ch^peus perpendicular in front, the nearly flat, verti- 
cal face extendmg upward to form a broad truncate horn about one- 
fourth wider than high. The face and sides of the horn reticulate, 
the enclosed spaces concave, shallow, and coarse; the truncate apex 
scarcely thickened, with the external angles very slightly nodose; 
the sides parallel over apical two-fifths, diverging posteriorly; margin 
thin, irregularly serrated, elevated just before the eye canthus to a 
very moderate right-angled prominence, the posterior edge of which 
continues posteriorly as a low, sharp carina separating the eye canthus 
from the flattened discal surface of the head. Canthus sloping away 
from the discal level of head; anterior and lateral margins sinuate to 



232990—53- 



104 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

the rounded, elevated, anterior angle, surface ridged inward from the 
angle, depressed along anterior edge and in posterior angle. Margin 
of head along inner edge of the eye with a reflexed carina, which grad- 
ually curves away from the eye, increases to a maximum height, then 
angles sharply inward and backward and sharply descends to general 
level of head ; head surface, including eye canthi, reticulate-tuberculate- 
punctate, very rough anteriorly on the concave posterior of the clypeal 
horn, gradually smoother over discal area, which is bounded poste- 
riorly by a low, rounded ridge between the eyes; the reticulation and 
tubercles of the discal area flatten until the outlines of the coarse, 
shallow punctures are barely traceable ; head behind the ridge smooth 
at middle but gradually roughened near the eyes. 

Pronotum similar to that of B. ferrugineus (Palisot de Beau vols) 
but with the horns of the median prominence and the high lateral 
edges of the depressions much farther forward. The anterior edge 
of the pronotum very finely margined, the middle third arching slightly 
over the base of the head, each outer third without a curve, straight, 
diverging anteriorly to the right-angled anterior angles, slightly 
concave close along these straight edges, then very obtusely convex 
and rounding into the deep lateral depressions, the middle third with 
a smaller, closer, parallel convexity and a moderate concavity at each 
end; pronotum laterally with two very large deep fossae on each side, 
the lower wider and less deep, widely sinuating the lateral margin 
from middle to near the anterior angles, the upper very deep, the 
outer edge thrown up in a high, thin, curving edge which descends 
to the posterior boundary of the lower cavity; posterior angles distinct 
but obtusely rounded; sides and base margined, the edge rather finely 
serrate laterally; pronotum convex basally; disc forming a broad 
median prominence with a sharp horn pointing upward on each side, 
the horns being rather sharply undercut below and basally flattened 
in front and behind; a deep median line from base forward over the 
median prominence and down between the horns. Surface of the 
pronotum alutaceous and finely scabrous back of the anterior margin, 
especially at the sides, then shallowly, coarsely punctate over the 
inner lower surface of the lateral fossae; posterior angles similarly 
finely alutaceous and closely, coarsely, shallowly punctate; elsewhere 
finely to minutely, sparsely punctate throughout, except for the very 
smooth surface of the deeper fossae; and with a large group of close 
very coarse punctures on each side back of the horn and above the 
deeper fossa, another group following the median line and spreading 
anteriorly to bases of the horns. 

scutellum finely punctate. Elytra not strongly shining; minutely 



BOLBOCERATIIsTI OF THE UNITED STATES — CARTWRIGHT 105 

alutaceous under high magnification; margined at base; striae mod- 
erately impressed, the strial punctures fine, moderately deep; first 
stria interrupted at scutellum; the second indistinct near scutellum. 

Metasternal lobe separating the middle coxae half as wide as the 
plate behind the coxae; lateral edges sharply carinate. 

Female unknown. 

Type.— Iloloiype male, USNM 61455. 

Type locality. — Sarita, Tex. Holotj^pe collected on bare sand, 
Nov. 30, 1911. 

Paratypes. — One male, CAS, Corpus Christi, Tex., June 28, 1942, 
E. S. Ross; one male, in Mark Robinson collection, Kingsville, Tex., 
C. T. Reed. 

Remarks. — Brady cinetulus rex may be readily separated from the 
two closely allied species B. ferrugineus (Palisot de Beauvois) and B. 
fossatus (Haldeman) by the obvious differences in the clypeal and 
thoracic horns. It may be noted also that the last tAvo species have 
only a single deep fossa on each side of the pronotum, while in B. rex 
this is doubled, the lower cavity sinuating the lateral margin. 

Bolbocerastes^ new genus 

This genus is closely allied to Bradycinetulus but differs in that the 
apex of the tibia of the midleg and hind leg is obliquely truncate, the 
scutellum is smooth or minutely punctate, the mandibles are parallel, 
nearly straight sided and bent sharply inward anteriorly, the sides and 
anterior edges forming a rectangle, the median prominence of the 
pronotum is without horns, two elevated carinae are present on each 
side of the pronotum, the prosternal spine behind the anterior coxae 
is transverse, doubly pointed and remote from the acutely angled 
intercoxal piece, and the aedeagus of the male is of a characteristic form 
(see fig. 14). 

Genotype. — Bolbocerastes regalis, new species. 

Key to the species of Bolbocerastes 

1. Anterior clypeal horn of male widely truncate; anterior and median horns 

of female usually equal in width, smooth back of median horn (Arizona, 

Nevada, California) regalis, new species 

Anterior clypeal horn of male narrowly truncate or rounded; anterior horn 
narrower than median horn in female, punctate back of median horn. 2 

2. Dull, dark reddish brown; elytral striae extremely fine, not impressed, the 

intervals completely flat even in reflected light, strial punctures very fine; 
fully developed male with the anterior clypeal horn narrowly rounded and 
very little advanced, much closer and more nearly in line with the strongly 
developed lateral horns than in serratus and imperialis (California, Baja 

California) peninsularis (SchaefFer) 

Elytral striae fine to moderate, usually at least slightly impressed, the intervals 
weakly convex in reflected light; fully developed males not as above 3 



106 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

3. Lateral serrations of pronotum arising between the two edges of a doubly 

carinate margin; elytral striae moderately impressed, strial punctures dis- 
tinct; color reddish brown, usually quite strongly shining; tip of clypeal 
horn rounded in male, inner thoracic carina without secondary angle at 
upper end; female cephahc horn rarely higher than clypeal horn, carina 
extending back from clypeal horn on each side sharp, strong, usually dark 
colored, lateral secondary horns nearer midway between clypeal and cephalic 
horns than to the cephalic horn, discal area of head distinctly punctate 
throughout, fine punctures of median thoracic prominence distinct and 
noticeable (Kansas, Oklahoma, Texas, New Mexico). serratus (LeConte) 
Pronotum without or with only traces of a doubly carinate margin laterally, 
the serrations appearing as thickenings of a single sharp edge; well developed 
males with a secondary angle at upper end of inner thoracic carina — 4 

4. Elytral striae much less distinct, the elytra smoother in appearance, nearly 

as in peninsularis , color lighter, more yellow-brown, less strongly shining; 
fully developed male with tip of clypeal horn narrowly truncate and well 
forward from the lateral horns, upper lateral thoracic carina of the male 
frequently with a noticeable secondary angle at upper end; female median 
cephalic horn often very high, usually higher than clypeal horn, carina 
extending back to lateral secondary horn weak, sometimes obhterated, not 
darker in color, the lateral secondary horns weak, usually much farther 
back, more nearly in line with the cephalic horn, discal area of head usually 
with only scattered punctures behind median horn (Baja California, Cali- 
fornia, Arizona, western Texas) imperialis, new species 

Elytral striae deeper, color red-brown, more strongly shining as in serratus, 
otherwise as above (Kansas) imperialis kansanus, new subspecies 

Bolbocerastes regalis, new species 

Figure 14, e; Plate 4 

Holotype male, length 18 mm., width 11.5 mm. In general appear- 
ance closely similar to Bolbocerastes serratus (LeConte), ferrugineus, 
shining, densely hairy beneath. Clypeus perpendicular in front, the 
flat vertical face extending upward to form a broad truncate horn, 
the vertical height subequal to base, the lateral edges arcuate from 
base to upper edge, narrowing at basal third to half the basal width 
then expanding to three-fourths, the upper edge weakly emarginate 
and subparallel with base, shining black in color, rounded and knoblike 
at each end; the sharply defined lateral triangular faces of the horn 
concave and posteriorly elevated from base into secondary horns half 
as high; posteriorly the broad curving slope of the clypeal horn merges 
into the flattened upper surface of the head, this quadrangular surface 
with a secondary horn at each anterior angle and with sharply defined 
carinalike subparallel lateral margins from which the prominent, 
widely explanate eye canthus slopes downward and outward. Occip- 
ital area smooth; gradually, finely, closely, shallowly punctate 
anteriorly; anterior and lateral faces of clypeal horn, labrum, eye 
canthi, and upper surface of mandibles finely reticulate-scabrous. 



BOLBOCERATESn OF THE UNITED STATES — CARTWRIGHT 107 

Broadly rounded labrum twice as wide as long. Upper surface of 
mandibles concave, sides straight, bent at right angles in front, 

Pronotum retuse in front; the deeper lateral impressions extending 
upward on each side of the median prominence and bounded laterally 
by two carinae on each side; the lower external carina starting near 
the serrate lateral thoracic margin and extending upward parallel 
with the basal thoracic margin for a distance equal to half the length 
of the lateral margin; the upper carina starting inside the upper end 
of the lower and extending upward for a slightly shorter distance; 
viewed from the side the lower carina is nearly straight, the upper 
end with a gentle downward curve, the lower end with a noticeable 
but low toothlike elevation; in outline the upper carina is half as 
high as long, sloping upward from a sharp basal curve to a rounded 
right-angled peak, then down on a concave curve to a noticeable sec- 
ondary angle. The median pronotal prominence sharply declivous in 
front, upper surface divided by an anteriorly increasingly depressed 
median line, the parallel sides with sharply rounded edges anteriorly; 
retuse front impunctate except for a subapical line of close, weak-to- 
obsolete punctures; median prominence with fine, close punctures 
shading to minute punctation across base; a few obsolete moderate 
punctures in the shallow lateral depressions outside the carinae; basal 
bead distinct. Lateral edges strongly serrate, as in Bolbocerastes 
serratus (LeConte). 

Elytra not strongly shining; striae scarcely impressed; strial 
punctures fine, moderately close, intervals practically flat, very 
minutely punctulate. 

Metasternal lobe separating middle coxae half as wide as plate 
behind the coxae. 

Female allotype, length 21 mm., width 10 mm. Similar to the male 
except that the thoracic prominences are much less developed and 
the head is different. The vertical face of the clypeus is only half as 
high as the basal length, the upper edge less than half the basal length, 
and from each end a curving carina extends outward to top of the 
secondary horn or tubercle; from the flattened area back of and about 
twice as high as the secondary tubercles arises a widely truncate 
median horn; anterior to this median horn the surface is reticulate- 
scabrous, posteriorly it is smooth. 

7V2)e.— Holotype male, USNM 61076. 

Type locality. — The holotype bears the following label data: "3206. 
Colo. R. bottom. Monument 204. Mex. Bd. line Mar. 20-31, '94. 
USNM Ace. No. 28133 Dr. E. A. Mearns." Monument 204 is approx- 
imately 20 miles south of Yuma, Ariz., near San Luis, Mexico, its 
exact location being lat. 32° 29' 03.59", long. 114° 46' 44.79". 



108 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

Allotype and paratypes. — Allotype and 97 paratypes bear data as 
follows: Nevada: 1, Hawthorn, June 8, 1943, Wallace; 1, Mount 
Charleston, 8,000-10,000 feet, Aug. 6, 1944, Zinn. 

California: 2, Baker, Mar. 27, 1935, and April 1935, Saylor; 1, 
California (State label only); 3, Coachella, May 12, 13, 17, 1927, Van 
Duzee; 3, Coachella, Riverside County, Apr. 28, 1928, May 9, 1927, 
July 15, 1927, Wymore; 2, Coachella Valley, May 9, 1927, Van Dyke, 
Aug. 20, 1949, V. d. Bosch; 1, Coyote Wells, Imperial County, June 5, 
1949, Hurd; 1, Essex, Apr. 29, 1937; 1, Holtville, June 28, 1936, 
Cazier; 14, Imperial County, Carrizo Light, Mar. 14, 17, 18, 22, 1928, 
Searl; 1, Indian Wells, Riverside County, Apr. 25, 1939, Martin; 
8, junction of Highways 78 and 99, Imperial County, Apr. 15, 1944, 
Tieman; 4, Mecca, Mar. 25, 1922, Benedict; 3, Murrays Dam, San 
Diego County, Cottle; 1, Needles, Apr. 1, 1941, Linsley; 2, Palm 
Springs, May 20, 1905, May 9, 1932; 2, San Clemente Island, Apr. 2, 3, 
1939, Channel Islands Biological Survey, Von Bloeker; 1, San Diego 
County, Mar. 10, 1928, light, Well; 7, Yermo, San Bernardino County, 
Apr. 11, 1936, Comstock, Sept. 6-30, 1939, Pierce. 

Arizona: 7, Arizona (State label only); 4, Ehrenberg, Oct. 18, 1938, 
Mar. 21, 23, 30, 1939, Parker; 6 (includes allotype), Gila Bend, Ma-. 1, 
1930, Lebart; 1, Holbrook, May 5, 1941, Dukes; 1, Navaho County, 
Duncan; 1, Phoenix; 1, Red Rock, Spring 1930, Burdett; 1, Tucson, 
April 192-, Duncan; 11, Yuma, February 1910, Carlson, Mar. 12, 18, 
1912, Slevin, May 15, 1939, Aitken. 

Mexico, Sonora: 1, Choya Bay, Mar. 27, 1949, Bradt; 1, 30 miles 
southwest of Sonoyta, 500 feet. Mar. 31, 1949, Bradt; 4, Puerto 
Libertad, Mar. 15, 1939, Parker. 

Paratypes in AMNH, CAS, LAC, KAN, MCZ, CAL, CU, OSJ, 
MICH, INHS, USNM, and the private collections of Mark Robinson, 
Henry Howden, C. A. Frost, A. T. McClay, F. H. Parker, D. K. 
Duncan, and R. H. Arnett. 

Remarks. — Bolbocerastes regalis varies from 16 to 20 mm. in length 
and from 10 to 12.5 mm. in width. It is closely similar to B. serratus 
(LeConte) but the males differ noticeably in having a higher, wide, 
truncate clypeal horn, smoother, less punctate pronotum, and on each 
side a doubly angulate inner pronotal carina which is always higher 
in the middle. In B. serratus (LeConte) the anterior clypeal horn is 
bluntly rounded, the pronotal punctures are more evident throughout, 
and the inner lateral pronotal carina is not doubly angulate or higher 
in the middle. The truncated tops of the anterior and median horns 
are about equal in width in females of B. regalis and the whole anterior 
part of the head, including clypeal and cephalic horns, appears higher 
than the posterior part, like an elevated snout. The lateral horns are 
most prominent in this species. 



BOLBOCERATINI OF THE UNITED STATES CARTWRIGHT 109 

Bolbocerastes imperialis, new species 

Figure 14, g; Plate 4 

Holotype male, length 19 mm., width 11 mm. Very similar to 
Bolbocerastes serratus (LeConte) ; ferrugineus; moderately shining; 
hairy beneath. Vertical face of clypeal horn inclined forward about 
30 degrees from perpendicular, slightly convex when viewed from side, 
height seven-tenths width at base, width of the truncate apex slightly 
less than half the basal width, the apex arcuate posteriorly at 
each side; the posterior face of the horn concave from side to side 
between the wealdy carinate edges, which converge very slightly from 
apex to base. General level of the head slightly more than half the 
height of the clypeal horn above the wide, truncate labrum; this flat- 
tened area sloping upward posteriorly and bounded by a ridge or 
carina deeply arcuate posteriorly from the anterior edge of one eye to 
the other and continuing forward on each side to the tips of the weakly 
elevated, right-angled, secondary horns, then converging to the an- 
terior edge of the clypeal horn and forming the upper edge of the tri- 
angular lateral area at base of the horn; flattened area of head slightly 
depressed laterally above the prominent quadrangular eye canthus, 
which slopes arcuately downward, outward, and finally upward at the 
outer angles; entire surface in front of the curving posterior ridge 
densely, rouglily punctate, smooth back of the ridge. Mandibles 
concave dorsally; strongly angled front and back. 

Pronotum retuse in front and with serrate lateral margins as in B. 
serratus (LeConte) and B. regalis; the median prominence and lateral 
carinae nearly as in B. regalis but with the outer carina with the upper 
end curving slightly away and not quite parallel with the basal mar- 
gin when viewed from the side; the inner lateral carina, viewed from 
the side, with the lower end rising abruptly to a right angle, then 
sloping evenly upward to a right-angled peak at middle, then down 
in a strongly sinuate in-and-out curve; both carinae, viewed from 
directly in front, evenly arcuate, with the peak angle of the inner carina 
strongly bent inward; the median prominence slightly constricted 
toward the base, with the sides weakly angulate in front of the con- 
striction and the surface more broadly and less deeply concave from 
side to side than in B. regalis and evenly arcuate from front to back; a 
distinct median line evident basally; the apical and deeper lateral 
pronotal depressions extending upward between the lateral carinae; 
the median prominence impunctate except for half a dozen or so close, 
moderate punctures on each side at base of the vertical sinus; above 
these about twenty or so scattered fine punctures on the inner slope 
of the vertical sinus, and a very few fine punctures indicating a sub- 
apical line; median prominence and lateral basal area back of the 



110 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

carinae closely, finely punctate; smoother and more finely punctate 
across base at middle; groups of moderate punctures in the depressions 
back of the carinae and an irregular row of close moderate punctures 
just inside the basal bead. 

Scutellum closely, minutely punctate. Elytra one-sixth wider than 
long; extremely, finely alutaceous under moderate magnification; 
striae scarcely impressed; strial punctures fine, distant about four 
diameters; intervals with very minute scattered punctures. 

Metasternal lobe separating middle coxae one-third as wide as the 
plate behind the coxae. 

Allotype female, length 14 mm., width 9 mm. Similar to the male 
except that the pronotal prominences are less strongly developed and 
the head is different. The vertical face of the clypeus is one-third 
as high as basal length; its upper truncate edge also one-third the 
length of the base; the concave triangular sides of the clypeal horn 
with the upper edge about twice as long as the anterior edge; the 
wide truncate frontal horn is about three-fourths as high as wide and 
about four times as high as the low secondary lateral horns, or tubercles. 

7V^^.— Holotype male, USNM 61077. 

Type locality. — Imperial County, Calif., "on the Experiment Farm," 
June 1912, J. C. Bridwell. 

Allotype and paratypes. — Allotype female and 210 paratypes bear 
the following data: Texas: 1, Castolon, June 1, 1928, Bibby; 1, 
Tornilla Flat, Big Bond National Park, July 12, 1948, desert, 3,650 
feet, at light, Werner and Nutting; 2, Boquillas, Brewster County, 
July 7, 1948, C. and P. Vaurie; 1, Marathon, June 14, 1948, D. Rocke- 
feller Expedition, Cazier; 1, Reeves County, June 10, 1939, Robinson. 

New Mexico: 1, Artesia, July 29, 1937, D. J. and J. N. Knull; 1, 
White Sands, July 23, 1933, Benedict. 

Arizona: 4, Agua Fria, Aug. 6, 1917, Bequaert, June 26, 1938, 
Duncan; 2, Aquila, Maricopa County, Aug. 21, 22, 1927, CU lot 542, 
sub 330; 2, Arlington, July 25, 1948, Anderson; 15, Arizona (State 
label only); 5, Avondale Ranch, Agua Fria River, Aug. 7, 1917, 
Wheeler leg.; 4, Baboquivari Mountains, July 24, 1941, Beamer, 
July 24, 1941, Todd; 2, Buckeye, July 30, 31, 1935, at light, Christen- 
son; 1, Cashion, Maricopa County, Aug. 25, 1949, at light, Werner 
and Nutting; 5, Cibola, August 1911; 1, Continental, Pima County, 
July 29, 1948, Ball; 1, Coyote Mountains, Aug. 4, 7, 1916; 1, Dome, 
July 21, 1924, Van Duzee; 4, Ehrenberg, July 20, 1946, June 12, 
1943, Grassman, July 18, 1939, Stager, June 21, 1938, Parker; 10, 
Globe, Duncan, Sept. 8, 1930, July 11, 1931, July, August, Duncan 
and Parker; 7, Hope, Yuma County, Aug. 12, 1948, greasewood desert, 
1,400 feet, at light, Werner and Nutting; 1, Hot Springs, July 16 
(year not given), at light, MorrUl; 2, Huachuca Mountains; 2, Kits 



BOLBOCERATIIsTI OF THE UNITED STATES — CARTWRIGHT 111 

Peak, Rincon, Baboquivari Mountains, Aug. 1, 4, 1916, about 1,050 
feet; 1, Littlefield ; 1, Marinette, Aug. 4, 1918, Schiffel; 1, Olberg, 
Sept. 13, 1936, Crandall; 1, Palmerlee; 8, Phinax (sic), September 
and November 1921; 17, Phoenix, Sept. 12, 1935, Crandell, May 12, 
1934, Parker, May 29, 1922, Odell, Aug. 2, 1917, CU Biol. Expedition, 
May 17, 1944, Parker, Apr. 29, 1941, Parker, May 7, 1918, Bradley, 
Sept. 12, 1926; 1, Pima County, Sept. 1, 1925, Marsh; 1, Prescott; 2, 
Sacoton, Oilman; 1, SafFord, September (no date given), Duncan; 
2, San Bernardino Ranch, Cochise County, 3,600 feet, sycamore- 
willow in valley at light, July 25, 1949, Werner and Nutting, July 16, 
1949, light, Parker; 1, San Luis, Yuma County, Aug. 11, 1940; 1, 
Talklai, Pollock; 20, Tucson, July 21 (no year given) Hubbard and 
Schwarz, July 30, 1937, Parker, August 1912, Breitenbecker, Aug. 20, 
1932, Flock, August 1935, Wharton, October 1927, Hamilton (2,500 
feet), July 10, 1950, Aug. 8, 1950, Bradt, Oct. 14, 1919; 3, Wellton, 
June 13, 1939, at light, Stitt; 1, Welton, Aug. 9, 1917, Wheeler leg.; 
1, Yuma. 

California: 7, Blythe, May 19, 1942, Allen, Aug. 26, Hurd, April 
16, 1945, Constantine, May 19, 1943, Vargas, July 22, 1947, July 8, 
1947; 4, Blythe, Riverside County, July 24, 1947, Aug. 24, 1947, 
MacSwain; 2, California (State label only); 29, Coachella, Riverside 
County, Apr. 28, 1927, May 9, 12, 17, 28, 1927, Wymore, May 22, 
1928, Van Dyke, May 13, 1917, Van Duzee; 2, Coachella Valley, 
Sept. 8, 1929, Comstock, Aug. 20, 1949, v. d. Bosch; 1, El Centro, 
Aug. 28, 1948, light trap; 8, Holtville, June 1936, Ross, August (no 
date given), Craig, June 28, 1936, Cazier, June 27, 1936, July 15, 
1936; 16 (includes allotype). Imperial County, June 1912, on Experi- 
ment Farm, Bridwell; 4, Imperial County, June 1925, Field, July 1, 
1925, Gehring; 2, Indio, July 17, 1923, Benedict, May 23, 1919, 
Slevin; 2, Meloland, June 4, 7, 1949, v. d. Bosch; 1, Needles, June 
12, 1940, Barr; 1, Neighbors, Apr. 27, 1930, Hornung; 1, Palo Alto, 
July 28, 1916; 1, Vallecito, San Diego County, Sept. 15, 1945, at light, 
Martin; 2, Warners, June 18, 1925. 

Mexico: 3, 10 miles south of Catavina, Baja California, July 29, 
1938, Michelbacher and Ross. 

Paratypes in CAS, LAC, AMNH, CARN, INHS, CAL, OSU, CU, 
MICH, TEX, KAN, CDA, USNM, and the private collections of 
R. H. Arnett, D. K. Duncan, C. A. Frost, P. C. Grassman, Henry 
Howden, A. T. McClay, Gayle H. Nelson, F. H. Parker, Mark 
Robinson, and F. Werner. 

Remarks. — Males of B. imperialis vary from 11 to 19 mm. in length, 
and from 7 to 11 mm., in width. In lesser developed males the punc- 
tures become fewer and weaker over the posterior third inside the 
arcuate carina at back of the head, anteriorly they persist strongly; 



112 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i03 

the clypeal horn decreases in height and its apex becomes narrow, 
sometimes rounded, but usually evidently truncate, and the anterior 
face vertical or sloping slightly backward; the median pronotal promi- 
nence is usually not constricted or narrowed basally. 

B. imperialis is allied to B. serratus (LeConte), B. regalis Cartwright, 
and to B. peninsularis (Schaeffer), which I believe is a distinct species. 
Well-developed males of B. imperialis are separated from the other 
species by the anterior clypeal horn being relatively farther forward 
from the lateral horns, the triangle formed by the tips of the three 
horns approaching an equilateral triangle much more closely than in 
the other species. The apex of the clypeal horn is much narrower 
than in B. regalis, and usually truncate rather than rounded as in 
serratus and peninsularis; in fully developed males the inner lateral 
carina is doubly angulate as in regalis but has the high middle peak 
bent inward, while in specimens of serratus and peninsularis this carina 
is not higher or angulate at middle. The color of serratus and penin- 
sularis is dark red-brown, of imperialis, a more yellow brown. 

In Bolbocerastes peninsularis, represented by 13 specimens, 3 of 
them cotypes, the male clypeal horn is rounded as in B. serratus but is 
much farther back relative to the lateral horns, which are nearly as 
high as the clypeal horn. The female is more heavily punctate, as in 
serratus, and the area back of the median horn is much more noticeably 
punctate than in imperialis. 

Bolbocerastes imperialis kansanus, new subspecies 

Holotype male, length 17 mm., width 11 mm. Very similar to B. 
imperialis; moderately shining ; darker in color; hairy beneath. Anterior 
face of the narrowly truncate clypeal horn vertical ; lateral secondary 
horns more prominent; surface back of the clypeal horn with a notice- 
able swelling on each side and diagonally backward from the posterior 
ridges of the horn to a point even with the forward edge of the eye 
canthus, leaving between them a widening depressed area which 
merges with the general posterior level of the head; surface anterior 
to the arcuate ridge between the eyes more concave and much smoother 
than in typical imperialis ; occiput smooth; eye canthus with a swelling 
diagonally inward from outer anterior angle. 

Pronotum more coarsely punctate throughout; lateral edges of the 
median prominence more nearly cariniform and not constricted 
basally; median peak of upper lateral carina not bent inward: the 
depression back of the anterior angles at bottom of the lateral sinuses 
deeper, more noticeable, and distinct; serrations of the lateral margin 
appearing as thickenings of a single sharp edge; otherwise similar to 
typical imperialis. 

Elytral striae deeper, nearly as in B. serratus (LeConte) ; the second 
stria basally indistinct over twice the length of the scutellum. 



BOLBOCERATINI OF THE UNITED STATES — CARTWRIGHT 113 

Aedeagus as in imperialis. 

Allotype female, length 16 mm., width 11 mm. Similar to typical 
B. imperialis except that punctm-es are coarser throughout; foveae 
are deeper; elytral striae are deeper; and the eye can thus has a convex 
swelling diagonally inward from the anterior angle. 

Type. — Holotype male, in Canadian Department of Agriculture 
collection. 

Type locality. — Kush County, Kan. 

Allotype and paratypes. — Allotype and 6 paratypes bear data as 
follows: Kansas: 4 (including allotype female), Eush County, Aug. 5, 
1920; 2, Ness County, Aug. 5, 1920; 1, Ness County, 2,260 feet, 
July 5, 1912, Williams. Paratypes vary from 13 to 17 mm. in length, 
and 8 to 11 mm. in width. 

Paratypes in CD A, KAN, and USNM. 

Remarks. — Bolbocerastes imperialis kansanus may be separated from 
B. serratus (LeConte) by the doubly angulate upper lateral thoracic 
carina in well-developed males, by the diagonal convexity of the eye 
canthus, by the lateral serrations of the pronotum appearing as 
thickenings of a single sharp margin, and by the second elytral stria 
all but disappearing at twice the length of the scutellum. In serratus 
the serrations of the pronotal margin arise between the two edges of 
the doubly carinate margin and the second elytral stria becomes 
indistinct opposite the apex of the scutellum. 

Bolbocerastes serratus (LeConte), new combination 

Figure 14, /; Plate 4 

Alhyreus serratus LeConte, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 80, 1854. 
Amechanus serratus Horn, Trans. Amer. Ent. Soc, vol. 3, p. 48, 1870. 
Bradycinetus serratus Horn, Proc. California Acad. Sci., ser. 2, vol. 4, p. 334, 1894. 
Athyreus (Bradicinetus) serratus Boucomont, in Wytsmann, Genera insectorum, 

fasc. 7, p. 8, 1902. 
Bradycinetus serratus Schaeffer, Trans. Amer. Ent. Soc, vol. 32, p. 251, 1906. 
Bradycinetulus serratus Cockerell, Ent. News, vol. 17, p. 242, 1906. 
Bolboceras (Amechamus) serratus Boucomont, Ann. Soc. Ent. France, vol. 79, 

p. 341, 1910. 

Bolbocerastes serratus (LeConte) and B. imperialis Cartwright, ex- 
cept for fully developed males, are frequently difficult to separate. 
The locality label will help in most cases since serratus is found east of 
New Mexico, while imperialis is found in California, Arizona, and 
rarely into west Texas. The maps (figs. 15 and 16) show the known 
distribution. In doubtful cases the male genitalia will separate the 
two (see fig. 14,/ and g). In serratus, the basal side or fl^oor of the 
cavity is formed by the two sides approaching each other with knife- 
edges, sharply angulate near the outside margin (in Bolbocerastes the 
aedeagus of the male is hooded apically by the two sides coming together 



114 



PROCEEDINGS OF THE NATIONAL MUSEUM 




• Bolbocerastes imperialis Cortwright x Bradyclnetulus rex Cartwright 

o Bradyclnetulus fossotus (Holdeman) 



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• Bolborhombus schoefferi (Boucomont) o Bolbelosmus orcuatus (Bates) 
X Bolbocerastes penlnsularis (Schoeffer) 

Figure 15. — Distribution of species of Bradyclnetulus and close'y related species. 



BOLBOCERATINI OF THE UNITED STATES — CARTWRIGHT 115 




e Bolbelasinus horni (Rivers) 
M Bolbelosmut minor (Linell) 



• Bradycinetulus ferruginous (Palisot de Beouvois) 

♦ Bolbocerastes Imperialis Itansanus Cartwright 




• Bolbocerostes regalls Cortwrlght o Bolborhombus ongulus (Robinson) 

• Bolbocerostes serrotus (Leconte) x Bolborhombus porvulus Cortwrlght 

Figure 16. — Distribution of species of Bradycinetulus'^&nd closely related species. 



116 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

to form a deep cavity) ; in imperiaJis, these edges are rounded and lead 
straight in without a break or angle. 

Type. — In LeConte collection, MCZ, 

Type locality. — Mexican Boundary, Laredo to Ringgold Barracks. 

Specimens examined. — 181. 

Distribution. — Kansas: Clark County. Oklahoma: Taloga, Chey- 
enne, *Alva, and *Payne County. Texas: Brownsville, *Beeville, 
Burnet County, Cypress Mills, Comal County, Cameron County, 
Caterina, Colorado City, Cotulla, Childress, Dimit County, Dallas, 
Dallas County, Edinburg, Eagle Pass, Fedor, Forestburg, Frio State 
Park, Goldthwaite, Hidalgo County, Kingsville, Laredo, Memphis, 
Menard, *Mineral Wells, New Braunsfels, Oakville, Plesanton, Reeves 
County, Round Mountain, Sabinal, Taloga, Uvalde, Weslaco, and 
Zavalla County. New Mexico: State label only (H. W. Wenzel col- 
lection). Mexico: La Gloria, south of Monclova, Coahuila; and 
Rancho Preso, Nuevo Le6n. 

Season. — April 28 to September 25, with largest numbers collected 
in May, June, and September. 

Bolbocerastes peninsularis (Schaeflfer), new combination 

FiQDRE 14, d; Plate 4 

Bradycinetus serratus var. peninsularis Schaeffer, Trans. Amer. Ent. Soc, vol. 32, 

p. 252, 1906. 
Bolboceras serratus var. peninsularis Boucomont, in Schenkling, Coleopterorum 

catalogus, pt. 46, p. 13, 1912. 

All specimens of this species examined have been dull, smooth, and 
of a uniform very dark red-brown color. The male is quite distinctive 
(see pi. 4, lower right). In the female the moderately high cephalic 
horn is twice the width of the tip of the clypeal horn and the sec- 
ondary lateral horns or tubercles are usually more prominent than in 
B. imperialis. The carinae between these and the clypeal horn are 
strong and well marked. 

Type. — Lectotype male, present selection, USNM 42568. 

Type locality. — Santa Rosa, Baja California, Mexico. 

Specimens examined. — 13. 

Distribution. — California: 2, State label only, CARN. Mexico: 
Santa Rosa, San Felipe, La Paz, and San Ignacio, all in Baja Cali- 
fornia. 

Season. — July 27 to October 1. 

Bolborhombus^ new genus 

Mandibles arcuate externally. Head, male and female, without 
median frontal horn. Pronotum serrate laterally; apex with a carina 
paralleling anterior margin, the interspace interrupted behind each 



I 



BOLBOCERATESri OF THE UNITED STATES — CARTWRIGHT 117 

eye by a distinct rather deep foveola; base margined. Scutellum tri- 
angular, with base straight, sides arcuate. Elytra not margined at 
base; seven discal striae, the first two interrupted by the scutellum, 
the second sometimes indistinctly forked opposite the apex of scutel- 
lum. Two prosternal spines in tandem behind anterior coxae, the 
posterior more or less hastate. Metasternal plate rhomboid in shape, 
the posterior angle acute and with adjacent edges cariniform, the 
lateral angles with adjacent edges rounded; the intercoxal lobe deeply 
concave and with strong cariniform lateral edges. 

Genotype.^ — Brady cinetus carinatus Schaeffer, 1906 = Bolboceras 
schaefferi Boucomont. 

Key to the species of Bolborhombus 

1. Clj'peus of male carinately margined in front, the very wide clypeal horn 

arcuate anteriorly, a low sharp carina arcuate in opposite direction behind 
the clypeal horn, which is midway between this and the carinate anterior 

margin; female unknown angulus (Robinson) 

Male clypeus not distinctly carinate along anterior edge but with parallel 
carinae extending back from anterior angles to the external angles of the tip 
of the horn 2 

2. Postapical carina of pronotum sharp and slightly undercut; male with short, 

distinct carina back of clypeal horn arcuate anteriorly, in same direction 
as clypeal horn; female smoother back of corresponding carina than in 
front of it; small shining species; 8 to 10 mm. (Baja California) . 

parvulus new species 
Postapical carina of pronotum rounded; male without carina or with faint 
indication of two low obscure tubercles back of clypeal horn; female equally 
rough behind and in front of carina; usually dull or only moderately shining 
(excepting some very large specimens, which may represent a subspecies) ; 
larger species, 11 to 18 mm schaefferi (Boucomont) 

Bolborhombus angulus (Robinson), new combination 

Figure 14, 6 

Bolboceras angulus Robinson, Trans. Amer. Ent. Soc, vol. 73, p. 170, 1947. 

Bolborhombus angulus (Robinson) is a small rufotestaceous species, 
7 to 11 mm. in length, readily recognized by the form of the head and 
clypeus. The wide straight anterior edge of the clypeus is strongly 
carinately margined. Behind this, from side to side and almost in 
line with the anterior edges of the ocular canthi, is a moderately high 
anteriorly arcuate ridge or horn, and still farther back a lower poste- 
riorly arcuate sharp carina. The median horn tends to become some- 
what nodose at the ends and is approximately midway between the 
posterior carina and the carinate anterior edge of the clypeus. In 
other characters angulus is very similar to B. parvulus and to small 
specimens of B. schaefferi (Boucomont). 

Type. — In Alark Robinson collection, Philadelphia, Pa. 



118 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Type locality. — Dog Canyon, Brewster County, Tex. 

Specimens examined.- — 4, including the holotype. 

Distribution. — Texas: Dog Canyon, Glenn Spring, and Boquillas, 
all in Brewster County. Arizona: Miller Canyon, Huachuca 
Mountains. 

Season. — July 7 to 29. 

Bolborhombus parvulus, new species 

Figure 14, a 

HolotjT^pe male, length 9 mm., width 6 mm. Shiny, rufotestaceous, 
densely hairy beneath. Clypeus extending obliquel}'^ upward and 
backward to top of clypeal horn; the anterior face practically flat, 
its surface closely, very coarsely punctate, with the forward edge 
convex and deeply reticulate-punctate, the lateral edges parallel and 
sharply carmiform; upper edge of the horn shallowly emarginate, the 
external angles only slightly tubcrculiform, from them a moderately 
high, sharp carina curving back each side and ending somewhat 
abruptly close to the innermost edge of the eye; surface of head back 
of horn uneven, with close, niLxed fine and coarse punctures and with a 
short, elevated, transverse, anteriorly arcuate carina midway between 
the similarly punctate, sharply edged, concave, depressed ocular canthi ; 
the carina about one-third as long as the width of the clypeal horn; 
the punctures back of the carina not quite as close as in front; the 
occiput smooth, closely, minutely punctate. 

Pronotum with three rather deep foveae close behind the finely 
margined anterior edge, one median, the others opposite mner eye 
margin on each side; a sharp postapical carina, slightly undercut on 
posterior side, starting on each side of the middle fovea and passing 
close behind the lateral foveae toward but not quite reaching the 
serrate side margin at about anterior third; a second short, similar 
carina on each side straight back from near the end of the postapical 
carina to the inner anterior edge of a strong lateral fovea; a shallow, 
coarsely punctate groove half way across the pronotum upward and 
backward from each of the tliree anterior foveae; a row of close coarse 
punctures in the lateral and basal marginal line, a few m the basal 
half of median line, and scattered coarse punctures on both sides of 
the lateral carma and continued upward to opposite the humerus, with 
mixed fine and minute punctation throughout; anterior angles slightly 
more than a right angle; posterior angles obtusely rounded; base 
distinctly margined. Scutellum with mLxed, very fine and minute 
punctation. 

Elytra not margined basally; striae weakly impressed; strial pimc- 
tures fine; first two striae interrupted by the scutellum; intervals 
weakly convex; the surface minutely punctate, very finely alutaceous 
under moderately high magnification. 



I 



BOLBOCERATIISri OF THE UNITED STATES — CARTWRIGHT 119 

Metasternal plate twice as wide as the lobe between the middle 
coxae, roughly punctate, and hairy along posterior edges, Prosternal 
spine 4-cornered, with the anterior angle curving do^vn to a sharp 
point between the coxae and with the lateral angles on a transverse 
level and the posterior curving do^vnward to form a sharp spine. 

Allotype female, length 9 mm., width 6 mm. Apparently identical 
with the male in every way except in characters of the head. Clypeus 
rising steeply at somewhat less than a right angle to a sharply carinate 
edge seven-eighths as wide and parallel to the turned-under anterior 
margin; the flat face enclosed being a little less than one-third as high 
as long, then dropping sharply and gradually up again with the 
lateral edges slightly diverging to the top of the very wide median 
horn, which is widely and deeply emarginate at middle, leaving the 
two ends, or external angles, as rounded, thickened, backward-sloping 
prominences; back of the emargination, on the same level, and at a 
point midway between the ocular canthi, is a very short arcuate 
carina or elongate tubercle, behind which the general level of the head 
is noticeably lower; surface in front of the horn deeply, coarsely 
punctate-reticulate, between the horn and the tubercle roughly 
coarsely punctate, and behind the tubercle much smoother, with 
dispersed, mixed fine and shallow, coarse punctures; occiput smooth, 
practically impunctate. 

Type.' — In California Academy of Sciences collection. 

Type locality.- — Triunfo, Baja California, Mexico. Holotype male 
collected Aug. 7, 1938, Michelbacher and Koss. 

Allotype and paratypes. — Allotype female, CAS, and paratype male, 
USNM 61656, San Venancio, Baja California, Mexico, Oct. 8, 1941, 
Ross and Bohart; paratype female, USNM 61656, Santa Rosa, Baja 
California, Mexico, Charles Schaeffer collection. 

Remarks.- — Bolborhombus parvulus is very near B. angulus (Robin- 
son) and is about the same size. The four specimens vary in length 
from 8)^ mm. for the smaller male to lOji mm. for the larger female. 
The anterior edge of the clypeus, described as margined in angulus, is 
almost exactly the same as in the female of parvulus. 

Bolborhombus schaefferi (Boucomont), new combination 

Figure 14, c 

Bradycinetus carinatus Schaeffer, Trans. Amer. Ent. Soc, vol. 32, p. 251, 1906. 
Bolboceras schaefferi Boucomont, Ann. Soc. Ent. France, vol. 79, p. 347, 1910. 

Bolborhombus schaeferi (Boucomont) averages much larger than 
B. parvulus and B. angulus, measuring 11 to 18 mm. in length, but it 
is very similar in all characters except those of the head, clypeus, and 
aedeagus. Males show considerable^variation in the clypeal horn. 



120 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i03 

In smaller specimens the clypeus slopes upward and backward from 
the wide straight anterior edge to the narrower but moderately wide 
and somewhat anteriorly arcuate apex of the horn, the external 
angles of which are thickened to slightly nodose. In large specimens 
the external angles of the horn become strongly elevated tubercles, in 
some individuals wide apart, in others relatively close together. The 
head back of the horn is densely, quite coarsely punctate, occasionally 
with a short, obscure, anteriorly arcuate carina or very low binodose 
tubercle. In the female the anterior edge of the clypeus is nearly 
vertical to the sharply carinate margin. Well back and nearly in line 
with the anterior edges of the ocular canthi are two widely separated, 
conical tubercles with a fine anteriorly angulate carina between them 
and a low median swelling or tubercle close behind them. The 
tubercles are usually strongly developed but occasionally in small 
specimens may appear only as slight thickenings of the ends of the 
angulate carina. 

Type. — ^Lectotype male, present selection, USNM 42569. 

Type locality. — Palmerlee, Cochise County, Ariz. 

Specimens examined. — 104. 

Distribution. — Arizona: Cave Creek and Pinery Canyon (Chira- 
cahua Mountains, Cochise County), Dewey, Douglas, Globe, Hua- 
chuca Mountains, Fort Huachuca, Kits Peak (Rincon, Baboquivari 
Mountains), Nogales, *Oracle, Palmerlee, Patagonia, Prescott, 
Reddington, Madera Canyon (Santa Rita Mountains), and Tucson. 
Texas: *Alpine, El Paso, *Fort Davis, Kingsville, and Round Moun- 
tain. Mexico: Alamos (Sonora), Venedio (Sinaloa), Oaxaca (Oaxaca), 
Santa Rosa (Baja California), Naco (Sonora), Tlahualilo, and Rio 
Mayo. 

Season. — July 1 to October 10, with largest numbers in July and 
August. 



B. S. 90VIRNMENT PRINTIH6 OFFICBi 1*13 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL. 103. PLATE 3 






•'"^f^rV 



^, 




Frcint and dorsal views of three species of Bradycinelulus: Left, B. fossatus (Haldeman); 
middle, B. rex, new species; right, B. ferrugineus (Palisot de Bcauvois). 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 103. PLATE 4 







Differences in the cl\'peai horn in Bolbocerastes: Upper left, B. serratus (LeConle); upper 
right, B. regalis, new species; lower left, B. impenalis, new species; lower right, B. 
peninsularis (Schaeffer). 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



byihe 




SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 



Vol. 103 Washington: 1953 No. 3319 

THE CHRYSOMELID BEETLES OF THE GENUS 
STRABALA CHEVROLAT 



By Doris Holmes Blake 



Introduction 



The first species of this chrysomehd genus was described by IlHger * 
in 1807 as Haltica ruja from Pennsylvania, Melsheimer collector. 
A year later, Olivier ^ described Altica ferruginea from Santo Do- 
mingo, Palisot de Beauvois collector, and Altica scutellaris from 
southern France. Of this latter locality von Harold ^ said that there 
was no Altica colored anything like this in southern France and that 
the locality was evidently an error. In 1837, in the Dejean catalogue, 
Chevrolat * listed under the new generic name Strabala {crpa^aKa, 
having a form thick and contracted) six specific names of Dejean 
without descriptions or references. Four of them are pure nomina 
nuda; for the two others, S. scutellata Dejean and S. dominicensis 
Dejean, the names {Altica) scutellaris? Olivier and (A.) ferruginea! 
Olivier, respectively, are given as doubtful synonyms. The generic 
name is consequently not published here. Chevrolat changed the 
locality for scutellata to America Borealis. Of these six names only 
one has since been used; S. intermedia, whose locality in this catalogue 
was given as Cuba, was adopted by Jacquelin Du Val ' in 1857 for a 

' Illiger, Mag. Insekt., vol. 6, p. 152, 1807. 

« Olivier, Entomologie, . . . vol. 6, pp. 697, 699, 1808. 

• von Harold, Ooleopterologische Hefte, vol. 14, p. 20, 1875. 

• Chevrolat, in Dejean, Catalogue de la collection de ColSopteres . . . ed. 3, p. 413, 1837. 

• Jacquelin Du Val, in Ramon de la Sagra, Historia . . . de la Isla de Cuba (Spanish ed.), vol. 7, p. IM, 
1857. 

232989—53 121 



122 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loe 

Cuban species. In d'Orbigny's Dictionnaire, Chevrolat' in 1848 
listed scutellaris and jerruginea under Strabala without the question 
marks, as well as the four nomina nuda he had given in 1837, thus 
establishing the genus. 

In 1868 Suffrian ' described Haltica amhulans from Cuba which he 
said belonged to Chevrolat's genus Strabala, a genus that Erichson ' 
had earlier merged with Lactica. Suffrian was not convinced, however, 
that these specimens having so poorly developed a basal sulcus on the 
prothorax really belonged to the genus Lactica. He stated that 
although there was an ill-defined basal sulcus, the beetles were smaller 
than most species of Altica and proportionately broader, and that the 
color was not blue but a deep reddish brown. This Cuban species that 
he had did not entirely correspond in coloring to Olivier's description 
of scutellaris or to Jacquelin Du Val's intermedia, both of which were 
paler beneath than the dark colored undersm-face of his beetles. 

Boheman ° described under the genus Strabala, two species, nigriceps 
from Buenos Aires and languida from Java, both of which have been 
referred to Lactica. Although I have not examined the types, the 
description of nigriceps as having the anterior angles of the prothorax 
truncate seems to exclude it from Strabala, and the description of 
languida is quite different from that of any of the group. Fall,^" in 
describing Altica testacea from Texas, compared it with rufa, noting 
that "rufa is much broader and less convex and has black legs and 
antennae." Fall's species, also, does not belong in Strabala. It differs 
in general shape, the prothorax has quite different anterior angles, 
and the scutellum is small and inconspicuous. 

Jacoby,^^ who had put scutellaris under Lactica, in the Supplement 
says of Horn, "Dr. Horn remarks that Lactica scutellaris would be 
better placed in the genus Haltica and that it is congeneric (and prob- 
ably also specifically identical) with Haltica rufa Uliger." But 
Jacoby "could not agree with this opinion" and kept scutellaris under 
Lactica. 

The British Museum (Natural History) specimens, therefore, are 
under Lactica, and in the U. S. National Museum, following Horn, 
they are under Altica. Henshaw ^^ lists rufa under Disonycha. 
Gemminger and Harold ^^ list rufa under Disonycha and scutellaris 
under Lactica. Heikertinger,'* in the Junk catalogue, nicely balances 

« OhevTolat, in d'Orbigny, Dictionnaire universe! d'histoire naturelle, vol. 12, p. 52, 1848. 
' Sufirian, Arch. Naturg., vol. 34, p. 182, 1868. 
» Erichson, Arch. Naturg., vol. 13, p. 173, 1847. 

• Boheman, Kongliga Svenska Fregatten Eugenies Resa . . . , vol. 2, Zoologi, pt. 1, Insects, pp. 189, 190, 
1850. 
M Fall, Trans. Amer. Ent. Soc, vol. 36, p. 157, 1910. 

" Jacoby, Biologia Oentrali-Americana, Coleoptera, vol. 6, pt. 1, p. 273, 1884; Supplement, p. 259, 1891. 
w Henshaw, List of the Coleoptera of America, north of Mexico, p. 112, 1885. 
>' Gemminger and von Harold, Oatalogus coleopterorum . . ., vol. 12, p. 3497, 1876. 
M Heikertinger, Coleopterorum catalogus, pars 166, pp. 241, 259, 1939. 



CHRYSOMELID GENUS STRABALA CHEVROLAT — BLAKE 123 

the dispute by synonymizing scutellaris and ambulans with rufa and 
placing them under Haltica and at the same time placing ferruginea 
and intermedia as separate species under Lactica. Mr. G. E. Bryant 
pointed out to me the specimens labeled Lactica scutellaris in the 
British Museum and said, "What are these doing here?" I agreed 
that it was high time to restore the original Chevrolat generic name 
Strabala for this group of very similarly colored species that no one 
has Itnown exactly what to do with. On my return to America, after 
a considerable search for the specimens of this species that had long 
been removed from the regular collection, I found that H. S. Barber 
had also labeled them Strabala. 

The species of this genus differ in color among themselves little more 
than do those of the genus Altica, but instead of being metallic blue 
they are a deep reddish or orange brown with dark or partly dark legs, 
dark scutellum (usually) , dark antennae, and more or less dark under- 
surface. Like Altica they have no spots, vittae, or other markings; 
and the variation in the dark coloration is slight. In practically all 
the North American specimens from Massachusetts to Panamd, the 
amount of dark coloration is very much the same, that is, the under- 
surface and femora are only partly dark. Hence the tendency has 
been to follow Horn/^ who wrote that ''Haltica rufa occurs from 
Massachusetts to Illinois, Florida and Texas, extending through 
Mexico to South America." There are, however, slight differences in 
the extent of the dark coloration, in the punctation, and in the shape 
of the aedeagus. In Eastern United States from New Hampshire to 
the Gulf and west to the Mississippi, Louisiana, and northern Texas, 
the species known as ruja Illiger presents little variation. It is a con- 
spicuously punctate, deep reddish brown species. In Florida occurs a 
race that, in dried specimens, at least, is always a pale yellow-brown, 
in contrast to the reddish color of rufa. It has also a distinctly aluta- 
ceous surface and is less distinctly punctate than rufa. In the speci- 
mens from Brownsville, Texas, to Panamd, which are shining and 
almost impunctate, in our limited collections are two groups with 
entirely differently shaped aedeagi — those that are narrowed at the 
tip and those with a wide, rounded tip. From the external appearance 
I cannot distinguish the beetles at all. H. S. Barber has labeled one 
lot, with round-tipped aedeagi, as a new species and the other as 
scutellaris. It seems to me that the name scutellaris cannot be applied 
to any species of Strabala with any certainty. To begin with, Olivier's 
locality of southern France, apparently erroneous, leaves us without 
any type locality in the western hemisphere. Species corresponding in 
coloration to Olivier's description occur in Cuba and Central America 
as well as North America. Therefore I propose to drop the name 

W Horn, Trans. Am. Ent. Soc, vol. 16, p. 232, 1889. 



124 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i03 

scutellaris entirely, as retaining it would only promote confusion. 
Among the specimens from Central America with the aedeagus nar- 
rowed at the tip there is a wide variation in the acuteness and in the 
width of the tip. I think there are probably a number of species, but 
since the coloring is so nearly uniform and the variation in the 
aedeagi only a matter of degree, I hesitate to give them specific names 
and am calling them only subspecies. 

In the Bowditch collection at the Museum of Comparative Zoology 
(MCZ) are seven specimens from South America (two from Venezuela 
and the rest from Colombia) that are uniformly somewhat larger than 
the others of the genus and, unlike the North and Central American 
specimens, are dark beneath, as in the majority of the West Indian 
specimens. In the British Museum is a series from Trinidad, col- 
lected by G. E. Bryant, also dark beneath but smaller, paler, and with 
a more rounded tip to the aedeagus. 

In the West Indies the prevailing color of the legs and undersurface 
of the specimens is entirely dark, but in one species, at least, the pos- 
terior femora are partly reddish, as in the continental species. In 
another species from Hispaniola, the scutellum is not black but 
reddish lil^e the elytra, the only one so colored that I have examined. 
Jacoby wrote of specimens with a red scuteUum from Juquila, Mexico, 
of which he had three. One specimen from Juquila in his collection 
is a yellow Lactica with a pronounced basal sulcus on the prothorax, 
typical of Lactica, and a pale scutellum. This may not be the one to 
which he referred. These slight differences in coloration of the speci- 
mens from the different islands of the West Indies are accompanied 
by a slightly different aedeagus. Still, as in the case of the Central 
American specimens, I hesitate to give them specific names and am 
calling them only subspecies. Much more work should be done on 
the group and many more specimens should be studied. Here I 
attempt to assemble the species that in the catalogues are divided 
among two or three genera and to call attention to their points of 
difference. 

The group is closer to Disonycha than to either Altica or Lactica. 
The head, while similar to Disonycha, has the circle of punctures 
nearer to the tubercles and the carina shorter and broader than in 
most species of Disonycha. The antennal joints are similar in being 
robust and rather short, but differ in having the third joint a little 
longer than the fourth, instead of the fourth longer than the third. 
The prothorax is much like that of Disonycha except that below the 
apical angle there is a tiny V-shaped notch which does not occur in 
Disonycha. The basal depression over the scuteUum, often found in 
Disonycha too, is longer and more pronounced in Strabala. The 
elytra are distinctive in being more convex and oval than in Lactica, 
Altica, or Disonycha. 



CHRYSOMELID GENUS STRABALA CHEVROLAT — BLAKE 125 




^.S.Qmbulans(5uffrloi) A.S.Qmbubos »sp JamalcsnatS ■. S ferru(JIoe<lK)iwier) y.S ambulons ssp. poerforicens(s 





k. Scolumblana 



/. S lnfcrmed{Q Jorq.duVat 



■. StrinltalO 



Figure 17. — Species of Strabala Chevrolat: a, rufa (Illiger); i, rufafloridana, new subspecies; 
c, acuminata, new species; d, rotunda, new species; e, acuminata teapensis, new sub- 
species /, acuminata costaricensis, new subspecies; g, ambulans (Suffrian); h, ambulant 
jamaicensis, new subspecies; i, ferruginea (Olivier); /, ambulans puertoricensis, new 
subspecies; k, columbiana, new species; /, intermedia Jacquelin Du Val; m, trinitatis, 
new species. 



126 PROCEEDESrGS OF THE NATIONAL MUSEUM vol. io3 

Genus Strabala Chevrolat 

Description. — Between 3.5 and 6 mm. in length, oval, deep yellowish 
or orange brown or reddish brown, with antennae dark, legs and under- 
surface more or less dark, and scutellum usually conspicuously dark; 
moderately shining, most species very minutely and confusedly 
punctate, only the North American species, iS. rufa (Illiger), very 
distinctly punctate. 

Head, except for mouth parts that are usually a little darker, deep 
yellowish or reddish, eyes entire and widely separated, the interocular 
space being more than half the width of the head; occiput smoothly 
rounded down to tubercles, with scattered and not very dense or coarse 
punctures; at the base of occiput, not visible except when the head is 
a little protruded, a groove running from behind one eye to the other; 
on either side above the frontal tubercles and near the eye a depression 
consisting of a large fovea, or more frequently a circle of punctures; 
frontal tubercles plainly marked, a broadly rounded and somewhat 
produced area between antennal sockets that extends and widens down 
the front nearly to the labrum, area under sockets a little depressed 
and on sides below eyes the cheeks rounded out and densely punctate. 
Antennae rather stout, not quite reaching the middle of the elytra, 
first two joints swollen, usually paler, remainder piceous, thh-d joint 
a little longer than fourth, fourth and fifth about equal. Prothorax 
not quite twice as wide as long, with rounded sides and narrow margin, 
the apical angle resembling that of Disonycha except that it is sepa- 
rated from the rest of the margin by a distinct though tiny V-shaped 
indentation, basal angle with a small tooth and obliquely cut, disk 
smooth and rather flat, with more or less of a depression along base 
and not distinctly limited at the ends as m Lactica. Scutellum large, 
usually conspicuously dark, contrasting with the reddish elytra. 
Elytra broader than prothorax, moderately convex, with small 
humeri and narrow margin, not depressed ; surface usually smooth, not 
distinctly punctate except in S. rufa, entirely yellowish or reddish 
brown. Epipleura broad, gradually diminishing and not quite reach- 
ing the apex. Body beneath variable in color, the prosternum red- 
dish, the breast and abdomen and legs in West Indian species usually 
but not always dark, in North American species the breast dark on 
the sides, the femora pale except in apical half or sometimes entirely 
pale, the tibiae and tarsi dark, the sides and apex of the abdomen 
sometimes also dark; in Trinidad and South American species, the 
coloring beneath dark as in the West Indian specimens. Anterior 
coxal cavities open, front coxae narrowly separated, legs rather short, 
hind femora moderately thickened, the tibiae as in Disonycha with a 
slight ridge, a spur at the end of posterior tibiae, claws appendiculate. 



CHRYSOMELID GENUS STRABALA CHEVROLAT — BLAKE 127 

Aedeagus a simple, curved structure with the tip varying in shape 
from acute to truncate and broadly rounded. 

Genotype. — Strahala ferruginea (Olivier), present designation. 

Key to the species of Strabala 

1. Scutellum reddish (Hispaniola) ferruginea (Olivier) 

Scutellum dark brown or piceous 2 

2. Species from the West Indies exclusive of Trinidad 3 

Species from North America 5 

Species from South America and Trinidad 9 

3. Body beneath more or less ferrugineous, hind femora ferrugineous at base, 

dark at apex (Cuba) intermedia Jacquelin Du Val 

Body beneath except prosternum more or less dark, hind femora entirely 
dark 4 

4. Aedeagus tapering gradually to a rounded tip (Cuba) ambulans (Suffrian) 

Aedeagus tapering gradually to a more acute but still rounded tip (Jamaica). 

ambulans jamaicensis, new subspecies 
Aedeagus distinctly truncate at tip (Puerto Rico). 

ambulans puertoricensis, new subspecies 

5. Elytra punctate or alutaceous or both (North of Mexico) 6 

Elytra not distinctly punctate or alutaceous (Texas and southward) 7 

6. Elytra deep reddish brown, distinctly punctate (New Hampshire south and 

westward to the Mississippi, Louisiana, Texas, and along the Gulf to Florida). 

rufa (lUiger) 

Elytra (in dried specimens) yellow, less distinctly punctate, alutaceous 

(Florida) rufa floridana, new subspecies 

7. Aedeagus well rounded at tip and with a slight constriction behind the tip 

(Mexico to Panamd) rotunda, new species 

Aedeagus gradually narrowing towards the tip, not constricted behind the 
tip 8 

8. Aedeagus narrowed to a rounded tip, hind femora dark at apex (Texas) . 

acuminata, new species 
Aedeagus narrowed to a truncate tip, hind femora entirely reddish brown 

( Mexico) acuminata teapensis, new subspecies 

Aedeagus narrowed to a broader truncate tip, hind femora varying from 
entirely reddish to reddish with a dark apex (Guatemala, Costa Rica) . 

acuminata costaricensis, new subspecies 

9. Large, 6 to 6 mm., aedeagus with an acute tip (Colombia, Venezuela). 

colombiana, new species 
Smaller, 4 to 5 mm., aedeagus with a somewhat truncate tip (Trinidad). 

trinitatis, new species 



Strabala rufa (Illiger) 

Figure 17, a 

Haliica rufa Illiger, Mag. Insekt., vol. 6, p. 152, 1807, 

Illiger's description of the thickly punctate elytra distinguishes 
this species from the others of the genus. The coloration is usually 



128 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

deep reddish brown above, with entirely dark antennae, dark scutel- 
lum, often entirely dark anterior legs, dark apical end of the posterior 
femora, dark breast, and sometimes also dark abdomen. Throughout 
its range over the country east of the Mississippi and in Louisiana and 
Texas, there seems to be very little variation except for the paler, 
less distinctly punctate race that occurs in Florida. The type locality 
is Pennsylvania, collected by Melsheimer. 

Distribution. — New Hampshire: Hampton, S. A. Shaw. Massachu- 
setts: Holyoke, Chicopee, F. Knab. Connecticut: Hartford, F. Knab. 
Rhode Island: Watch Hill, W. Robinson. New York: West Point, 
W. Robinson. New Jersey: Boynton. Illinois: C. Thomas. Mich- 
igan: Detroit, Hubbard and Schwarz. Maryland. Washington, 
D. C. Virginia: Fredericksburg, Clifton, J. C. Bridwell; Nelson 
County, W. Robinson; Stone Creek, Lee County, Hubbard and 
Schwarz; Vienna, J. C. Bridwell; West Falls Church, E. A. Chapin. 
West Virginia: Aurora. Kentucky: H.Soltau. Tennessee: Memphis, 
H. Soltau. North Carolina: Graybeard Mountain. South Carolina: 
Florence. Georgia: Savannah. Alabama: Coleta, H. M. Smith. 
Mississippi: Greenwood, T. H. Parks. Louisiana: Baton Rouge, 
T. H. Jones; Port Allen, J. L. Lauderdale; New Orleans, H. Soltau. 
Texas: Liberty, R. A. Vickery. Two specimens, both females, 
USNM (Brooklyn Museum collection), labeled simply "Mexico," are 
apparently this species. 

Strabala riifa floridana, new subspecies 

Figure 17, 6 

This race differs from the reddish bro\\Ti one found elsewhere in 
eastern United States by being (in dried specimens) pale yellow-brown 
with the dark markings as in typical rufa. The elytra are less dis- 
tinctly punctate than in rufa and distinctly alutaceous. 

Tyjpe. — Holotype male and 7 paratypes, USNM 61202, labeled 
simply "Fla.," collector F. Knab. 

Distribution. — Florida: Lake Ashby, Capron, Indian River, all 
collected by Hubbard and Schwarz; Lake Wales, Lakeland, E. M, 
Craighead; Jacksonville, Ashmead; Sanford, Van Duzee; St. Peters- 
burg (on sweet corn) ; Ft. Myers (on sweet and white potato) ; Oneco 
(on spinach); Homestead (on string beans); West Palm Beach (on 
sweet potato and lima beans) . 

Strabala acuminata, uew species 

Figure 17, c 

From 4.5 to 5.5 mm. in length, oval, shining, orange-brown, with 
dark antennae, dark scutellum, darkened sides to breast, and dark 
anterior legs and apical part of posterior femora, tibiae, and tarsi. 



CHRYSOMELID GEsSTUS STRABALA CHEVROLAT — BLAKE 129 

Head shiny, only a few scattered punctures over occiput besides 
the fovea or circle of depressed punctures near the eye; mouth parts 
frequently edged with deeper brown. Antennae deep brown, with 
the three basal joints frequently paler. Prothorax shiny, almost 
impunctate, only under high magnification are very fine punctures 
visible ; basal sulcus usually not very long or pronounced, but somewhat 
variable in different specimens, as is usual in the genus. Scutellum 
dark. Elytra mirror smooth, shining, impunctate, sometimes very 
indistinctly alutaceous. Body beneath with sides of breast, anterior 
legs, apical part of posterior femora, tibiae, and tarsi dark. Length 
4.4 to 5.5 mm.; width 2. 4 to 3 mm. 

Type.— Holotype male and 2 paratypes, USNM 61203 (Brooklyn 
Museum collection), collected at the Esperanza Ranch, Brownsville, 
Texas, August 18 (no year given). 

Other localities. — Texas: Brownsville, R. A. Vickery; another speci- 
men, from same locality (on string bean leaves); San Benito (on 
bean foliage); Hidalgo (on black-eyed-pea leaf). 

Remarks. — The shiny, almost impunctate surface with little evidence 
of alutaceous dullness at once differentiates this from Strabala rufa 
(lUiger). There are no records of it farther north than the Brownsville 
region. 

Strabala acuminata teapensis, new subspecies 

Figure 17, e 

Lactica scutellaris Olivier, Jacoby (part), Biologia Centrali- Americana, vol. 6, 
pt. 1, p. 273, 1884. 

This differs from the Brownsville, Texas, race in having less dark 
coloring, the femora all being deep reddish, the tibiae and tarsi only 
dark, and the sides of the breast not so dark. The upper surface is 
a shining deep reddish brown. The aedeagus has a somewhat trim- 
cate tip. 

Type. — Holotype male, in the British Museum (Natiu-al History), 
from the Biologia Centrali-Americana material, labeled by Jacoby 
Lactica scutellatis Olivier, from Teapa, Tabasco, Mexico, collected in 
March by H. H. Smith; 1 paratype in U. S. National Museum, 
USNM 61204; and 2 paratypes in Museum of Comparative Zoology 
from same series. 

Strabala acuminata costaricensis, new subspecies 

Figure 17, / 

Lactica scutellaris Olivier, Jacoby (part), Biologia Centrali-Americana, vol. 6, 
pt. 1, p. 273, 1884. 

This differs from the Brownsville, Texas, and Teapa, Mexico, races 
by having a still wider truncate tip to the aedeagus. In coloring, it 



130 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i03 

is somewhat deeper red than the Brownsville specimens and differs 
from the Teapa specimens in having the anterior femora and apical 
part of the posterior femora as well as the sides of the breast dark. 

Type. — Holotype male and 1 paratype, USNAl 61205, collected at 
Vara Blanca, Costa Rica, 1,700 meters, July 7, 1928, by F. Never- 
mann; 1 paratype in British Museum (Natural History). 

Other localities. — Costa Rica: San Jose, 1,000-2,000 meters, Aug. 25, 
1928, F. Nevermann. Guatemala: Yepocapa, August 1948, H. T. 
Dalmat; Cob^n, Alta Vera Paz, May 1926, J. M. Aldrich; Purula, 
Champion 

Strabala rotunda, new species 

Figure 17, d 

Lactica scutellaris Olivier, Jacoby (part), Biologia Centrali- Americana, vol. 6, 
pt. 1, p. 273, 1884. 

From 4 to 5.5 mm. in length, oval, shining although faintly alu- 
taceous, very finely punctate, yellowish or reddish brown with dark 
antennae, dark apex to femora, dark tibiae and tarsi, and the breast 
and abdomen often darkened on the sides, varying to entirely dark 
(rare) . 

Head reddish, with the mouth parts usually deeper brown; finely 
punctate over occiput, with the usual fovea or circle of depressed 
punctures. Pro thorax shiny and very finely punctate with a sulcus 
along the base well developed. Scutellum dark. Elytra faintly 
alutaceous and finely punctate. Body beneath varying in color 
from having the sides of breast and abdomen deeper brown to being 
entirely dark, the latter rare ; femora with apical half dark, tibiae and 
tarsi dark. Length 4 to 5.4 mm.; width 2.6 to 3 mm. 

Type. — Holotype male, USNM 61206, and 3 paratypes in British 
Museum (Natural History) collected at Volc^n de ChiriquI, Panamd, 
by Champion. 

Other localities. — Panamd: Panama, Aug. 13, 1946, by N. L. H. 
Kxauss; Almirante, Sept. 4, 1938, H. Dybas; Bella Vista, Aug. 7, 1924, 
N. Banks; Mindi Dairy (on para grass), Mindi, Canal Zone, July 
1918, H. F. Dietz and J. Zetek; Plantation Borracho, Canal Zone 
(swept from grass under papaya trees), July 1918, H. F. Dietz and 
J. Zetek; Bugaba, Champion (in British Museum). Guatemala: 
Trece Aguas, Alta Vera Paz, May (date not given), H. S. Barber and 
E. A. Schwarz. Mexico: In bananas from Mexico taken at Mobile, 
Alabama, July 19, 1929; with orchids from Guerrero (State), taken at 
Laredo, Texas, Feb. 28, 1940; with orchids from Tamazunchale, 
San Luis Potosi, taken at Laredo, Texas, Feb. 28, 1940; with tomatoes 
from El Mante, Tamaulipas, taken at El Paso, Aug. 1, 1947; Colima, 



CHRYSOMELID GENUS STRABALA CHEVROLAT BLAKE 131 

Hoge and Conradt; Jalapa, Hoge; on green peppers from Manuel, 
Tamaulipas, taken at Laredo, Texas, Jan. 7, 1935. 

Remarks. — H. S. Barber has attached a specific name from the 
Mexican locaUties to this species, but since the range is as far south 
as Panamd, his name would be somewhat misleading. The rounding 
of the tip of the aedeagus varies considerably but the slight con- 
striction behind the tip is always present, and sets the species apart 
from the rest. The only other species with a similar aedeagus tip 
in the group is Strabala intermedia Jacquelin Du Val, from Cuba. 
In all, 15 specimens of S. rotunda have been taken on orchids 
from Mexico at the port of entry to this country. 

Strabala ambulans (SufiFrian) 

Figure 17, g 
Haltica ambulans Sufifrian, Arch. Naturg., vol. 34, p. 182, 1868. 

Specimens from Cuba agree with SufFrian's detailed description of 
his Cuban H. ambulans. The upper surface is faintly alutaceous and 
finely punctate, in color a deep reddish brown with dark scutellum 
and dark legs, breast, and abdomen. 

Distribution. — Cuba: Cayamas, E. A. Schwarz; Central Jaronu, 
L. C. Scaramuzza; Baragua, L. C. Scaramuzza, L. D. Chiistenson 
(MCZ); upper Yara Valley, Oct. 18, 1928, L. C. Scaramuzza (MCZ) ; 
Soledad, Cienfuegos, Nov. 7, 1926, P. J. Darlington (MCZ); Aguada 
(MCZ); Mountains north of Imias, eastern Oriente Province, July 
25-28, 3-4,000 feet, P. J. Darhngton (MCZ). 

Strabala ambulans jamaicensis, new subspecies 

Figure 17, h 

Between 3.5 and 4,5 mm. in length, oval, faintly shining, indis- 
tinctly alutaceous, very finely punctate, reddish brown, the mouth 
parts, antennae, legs, breast, and abdomen dark. 

Head with scattered punctures, the usual fovea or circle of punc- 
tures between eye and frontal tubercle on each side. Antennae with 
the basal joints a little paler. Prothorax having a distinct sulcus 
along the base. Scutellum dark. Elytra very faintly punctate. 
Body beneath with breast, abdomen, and legs dark. Length 3.5 to 
4.5 mm.; width 2 to 2.3 mm. 

Type. — Holotype male and 10 paratypes, USNM 61207, collected 
at Manchioneal, Jamaica, Jan. 30, 1939, by E. A. Chapin and R. E, 
Blackwelder. 

Other localities. — Jamaica: Roaring River Falls, St. Ann's Parish, 
Sept. 13, 1917, H. Morrison; Fern Gully, St. Ann's Parish, Sept. 14, 



132 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

1917, H. Morrison; Mandeville, Van Duzee, A. E. Wight (MCZ); 
Montego Bay, Mar. 2, 1911; Balaclava (MCZ). 

Remarks. — This subspecies differs only slightly from the Cuban race 
in having a more acute though still rounded tip to the aedeagus. 

Strabala ambulans puertoricensis, new subspecies 

Figure 17, j 

Between 4.5 and 5 mm. in length, oval, shiny, elytra not alutaceous, 
very indistinctly punctate, reddish brown, antennae, mouth parts, 
legs, breast and abdomen dark. 

Head with scattered punctures and a fovea or circle of punctures 
near the eyes. Antennae with paler basal joints. Prothorax not at 
all clearly depressed along the base. Scutellum dark. Elytra 
quite shiny, not alutaceous, very finely and indistinctly punctate. 
Body beneath dark except the prosternum. Length 4.5 to 5 mm.; 
width 2.5 to 3 mm. 

Type. — Holotype male and 1 paratype, USNM 61208; 1 paratype 
in Museum of Comparative Zoology, taken at Adjuntas, Puerto Rico, 
Nov. 1, 1932, by R. G. Oakley. 

Other localities. — Puerto Rico: Lares, September 1921, G. N. Wol- 
cott; Arecibo, June 17, 1932 (on eggplant leaf), Anderson and Mills; 
Mayaguez, January 1899, A. Busck; Villalba, June 18, 1934, C. M. 
Matos (MCZ); Mayaguez, December 1932, V. Alexandrina (MCZ). 

Remarks. — The shiny, not alutaceous elytra, the larger size, and the 
broad, almost truncate tip of the aedeagus are unlike the Cuban and 
Jamaican races of the species. 

Strabala ferruginea (Olivier) 

Figure 17, i 

Altica ferruginea Olivier, Entomologie . . . , vol. 6, p. 697, 1808. 
Strabala ferruginea Chevrolat, in d'Orbigny, Dictionnaire universel d'histoire 
naturelle, vol. 12, p. 52, 1848. 

Olivier's description of the scutellum of this species as ferrugineous 
would differentiate it from the rest of the West Indian species even if 
he had not given the type locality as Santo Domingo. He described 
the color of the undersurface as black and this agrees with many of 
the specimens, but there is a certain variation in the coloring, some 
specimens being entirely ferrugineous beneath, others with dark legs 
and reddish breast and abdomen, others with dark legs, breast, and 
abdomen. Moreover the pronotum and elytra are finely and a little 
more distinctly punctate than in Cuban and Jamaican species and 
not at all alutaceous. The aedeagus, having a truncate tip, resembles 
the Puerto Rican specimens more than the Cuban or Jamaican. 



CHRYSOMELID GENUS STRABALA CHEVROLAT — BLAKE 133 

Distribution. — Haiti: La Vanneau, June 1920, Bizotan, Nov. 3, 
1925, Diguini, Apr. 14, 1925, all collected by W. A. Hoffman; Grande 
Riviere, W. M. Mann; He de la Tortue, April 1929, E. C. and G. M. 
Leonard; Port-au-Prince and vicinity, October 1934, P. J. Darlington 
(MCZ); Miragoane, October, November, P. J. Darlington (MCZ); 
Carrefour, May 1-3, 1908, M. Cameron (British Museum). Dominican 
Republic: August Busck; Macoris, Mar. 26, 1913, P. G, Russell; Puerto 
Plata, Hurst (MCZ); Santiago, 1938, P. J. Darlington (MCZ) 
Slinchez, July 1938, P. J. Darlington (MCZ). 

Strabala intermedia Jacquelin Du Val 

Figure 17, I 

Strabala intermedia Jacquelin Du Val, in Ram6n de la Sagra, Historia . . . de la 
Isla de Cuba (Spanish ed.), vol. 7, p. 129, 1857. 

In coloring, three old specimens in the Museum of Comparative 
Zoology labeled "Cuba" resemble Jacquelin Du Val's short description. 
They are deep yellowish brown with dark antennae, dark scutellum, 
dark tibiae and tarsi; the anterior femora and the apex of the pos- 
terior femora are dark. In size they are slightly larger than the 
measurements given for S. intermedia; instead of being 3% mm. they 
are 4.5 to 5.5 mm. in length. In two, both females, the basal thoracic 
sulcus is pronounced, almost like that of a true Lactica, but in the third 
specimen, a male, the depression along the base is less marked. The 
surface is faintly alutaceous and iBnely punctate. In aU three speci- 
mens the head is rather densely punctate in addition to the circle 
of depressed punctures near the eye. The coloring of these specimens 
is more like that of the North American species. The aedeagus, 
although essentially the same as in all the others, is distinctive in 
being slightly constricted behind the tip as in the Mexican and Central 
American species S. rotunda, but differs in having a more acute tip. 

Strabala colomhiana, new species 

Figure 17, k 

From 5 to 6 mm. in length, oblong oval, deep reddish brown, the 
antennae, breast, abdomen, and legs dark, shining, not alutaceous, 
very finely, and the elytra rather densely, punctate. 

Head usually entirely deep reddish brown, shining, with scattered 
fine punctures over occiput and the usual depressed circle of punctiu-es 
or fovea near eye. Antennae short, dark, the two basal joints paler. 
Prothorax shiny, finely punctate, a depressed line near base. Scutel- 
lum dark. Elytra shiny, very finely and indistinctly punctate. Body 
beneath with breast and abdomen deep piceous, legs dark. Length 
5 to 6 mm.; width 2.8 to 3.2 mm. 



134 PROCEEDINGS OF THE NATIONAL MUSEUM 



Type. — Holotype male, in Museum of Comparative Zoology, and 
1 para type, USNM 61209, from Bogota, Colombia, from the Bow- 
ditch collection, Jacoby's second collection. 

Other localities. — Colombia; Nueva Granada; Venezuela; and Cara- 
cas; all in the Bowditch collection, Jacoby's second collection. 

Remarks. — Among the material presented to the U. S. National 
Museum from the Bowditch collection is a single old specimen 
of Strabala sp. with the label "Lactica—Strabala haematina Dej" 
from "Nov. Granad." This may be one of the specimens listed in 
the Dejean Catalogue from Cartagena since it bears the old Dejean 
Catalogue Dame. The aedeagus of this old specimen has a rounded 
tip quite unlike the one from Bogota with its acute tip, and is probably 
not the same species. 

Strabala trinitatis, new species 

Figure 17, m 

From 4 to 4.9 mm. in length, oval, shiny, not alutaceous, finely 
but distinctly punctate, yellowish or reddish brown with darker 
antennae, scutellum, legs, breast and abdomen. 

Head entirely reddish, shiny, finely punctate over occiput. Anten- 
nae dark, with the first two or three joints paler. Prothorax shiny, 
finely punctate, with a moderately developed sulcus over the scutel- 
lum. Elytra distinctly and rather densely punctate, shiny, not at 
all alutaceous. Body beneath with breast, abdomen, and legs deep 
reddish brown. Length 4 to 4.9 mm.; width 2.5 to 2.7 mm. 

Type. — Holotype and 4 paratypes, British Museum, and 1 paratype, 
USNM 61210, collected m Trinidad m 1903 by G. E. Bryant. 

Other localities. — Two specimens (USNM) collected by A. Busck on 
June 12 at Montserrat, Trinidad. 

Remarks. — The specimens in this series, although of the coloration of 
the ones from Colombia and Venezuela, are smaller, more oval, and 
distinctly though finely punctate. The tip of the aedeagus is truncate 
and not at all acute as in the single male specimen from Bogota. 



B. S. GOVERNHENT PRINTING OFPICEi I9SS 




PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



Issued 



SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 
Vol. 103 Washington: 1953 No. 3320 

AMERICAN BITING MIDGES OF THE HELEID GENUS 

MONOHELEA 



By Willis W. Wirth* 



The genus Monohelea was founded by Eaeffer (1917) for a new 
species, hieroglyphica, from Paraguay, and three other American 
species were included: Ceratopogon seguax Williston, 1896; C. macuLi- 
pennis CoquLllett, 1905; and C. nebulosa Coquillett, 1901. Since then 
a number of species have been added to the genus, including repre- 
sentatives from each of the world's zoogeographic regions. However, 
with the exception of Lee's (1948) treatment of the eight Australasian 
species, no comprehensive paper on the genus has yet appeared. 

A study of the American Monohelea has resulted in the recognition 
of 7 undescribed species, bringing the total for the hemisphere to 13. 
No doubt many more await discovery. These American species fall 
in four distinct groups, each with characteristic wing and leg markings. 
Within these groups the species are rather difficult to separate, for the 
characters which prove useful in one group may be valueless in others. 
These groups may also be recognized by characters of the male geni- 
talia, which offer in addition good specific characters. 

The generic position of Ceratopogon seguax Williston, based on a 
male from St. Vincent, cannot be determined with certainty until 
the male genitalia or the female are studied. According to the original 
description, seguax would probably fit as well in Stilobezzia. Simi- 
larly, the male recorded as Monohelea sp. by Floch and Abonnenc 
(1942) from French Guiana could be either Monohelea or Stilobezzia, 
since in their figure of the male genitalia the aedeagus is omitted. 

The types of five of the new species, as well as most of the specimens 
studied, are in the U. S. National Museum. For the loan or donation 

' Bureau of Entomology and Plant? Quarantine, Agricultural Research Administration, U. S. Depart- 
ment of Agriculture. 

232992—53 135 



136 PROCEEDINGS OF THE NATIONAL MUSEUM ^°^- "^ 

of other specimens examined I am deeply grateful to Paul Freeman, 
of the British Museum (Natural History), to Henry Dietrich, of 
Cornell University, and to John Lane, of the University of Sao Paulo. 
For the generous gift or loan of still other specimens I wish to thank 
Mrs. Elisabeth C. Beck and J. A, Mulrennan of Jacksonville, Florida, 
Bernard Brookman of San Francisco, California, Jean A. Laffoon of 
Ames, Iowa, and Henry K. Townes of Raleigh, North Carolina. 
The figures of the wings and legs were made with the aid of a micro- 
projector and those of the male genitalia with the aid of an ocular 
reticule disc and squared paper. No consistent scale was used. 

Family Heleidae 

Genus Monohelea Kieffer, 1917 

Diagnosis. — Body short, not very haiiy. Eyes bare. Mesonotal 
pits present but small. Forelegs and midlegs umnodified; fourth 
tarsal segments cylindrical and claws small and equal in both sexes; 
hind leg with femur and tibia more or less thickened, but without 
spines, basitarsus with a spine at tip, fourth segment cylindrical, fifth 
segment in female with one very long claw, the other absent or very 
short, male claws small and equal except hind claw as in female in 
tessellata group; empodium absent. Wings with fine micro trichiae, a 
few macrotrichiae at wing tip ; costa extending well beyond middle of 
wing; two radial cells, second longer than first; intercalary fork fairly 
distinct; crossvein r-m vertical; median fork with short stem, M2 
sometimes interrupted at base; anal vein thickened in middle, but 
without fold at thickening. Male genitalia with ninth tergite tapered, 
the caudal margin truncate or broadly bilobed; aedeagus usually with 
a pair of sharp-pointed, tapered, lateral sclerites and an accessory, 
dorsomedian, posterior lobe; parameres consisting of paired, sub- 
median sclerites of irregular shapes. 

Key to the American species of Monohelea 

1. Wings without markings; mesonotum shining black (subgenus Schizohelea) . 

1. leucopeza (Meigen) 
Wings with pronounced dark markings; mesonotum pruinose, often with mark- 
ings (subgenus Monohelea) 2 

2. Hind femur and tibia yellow, with dark brown rings {hieroglyphica group) . 3 
Hind femur and tibia shining black, only knee spot yellow 9 

3. Forelegs and midlegs with distinct dark brown bands; wings often with exten- 

sive macrotrichiae 4 

Forelegs and midlegs yellowish, without distinct bands; wings with macro- 
trichiae only at extreme distal margin 6 

4. Wings with macrotrichiae over distal third; wing markings extensive 5 

Wing without apparent macrotrichiae; wing markings reduced, X-shaped 

marking in cells Mi and M2 broken up into 3 discrete spots. 

6. texana, new species 



HELEID MIDGES, GENUS MONOHELEA — ^WIRTH 137 

5. Wing with about 20 small, scattered, black dots in addition to the irregular, 

diffuse gray markings; fore tibia and midtibia dark only at apices; scutellum 

uniformly whitish 8. brasiliensis Lane 

Wing with uniformly gray, irregular markings, without small, black dots; 
fore tibia and midtibia entirely dark; scutellum dark in middle. 

7. ornata, new species 

6. Hind femur brown on basal third; scutellum dark in middle; wing markings 

not whitish bordered 7 

Hind femur with broad yellow band on sub-basal fourth; scutellum uniformly 
dull whitish; wing markings bordered with whitish pruinosity. 

3. maculipennis (Coquillett) 

7. Basal and median brown bands on hind tibia joined with brown, at least on 

ventral side ; male parameres each with 2 or 3 distal lobes 8 

Basal and median brown tibial bands separated by a broad yellow band ; male 
parameres each with simple, bladelike apex 5. macfiei, new species 

8. Sinuate, dark, wing marking in cells Mj and M2 entire; male parameres with 

broad, rounded distal lobes 4. lanei, new species 

Sinuate, dark, wing marking broken into 3 or 4 separate spots; male parameres 
with slender, pointed, distal lobes 2. hieroglyphica Kieffer 

9. Scutellum yellow, brown in middle; wing with 2 prominent, small, black spots, 

spot absent in basal cell; hind basitarsus about 2.5 times as long as second 
segment ; mesonotum yellowish, with tiny brown dots (multilineata group) 10 
Scutellum entirely dark brown or yellow only in middle ; wing with two prom- 
inent anterior spots plus a spot in basal cell; hind basitarsus 1.3 to 2.0 times 
as long as second segment {tessellata group) 12 

10. Ninth sternite of male with 4 long hairs arising from tubercles in a row at base 

of median lobe 11 

Ninth sternite of male with 2 long hairs arising from base of median lobe; para- 
meres not connected, with irregular, twisted apices; female abdomen 
pruinose gray above, with small, lateral, shining brown spots. 

9. stonei, new species 

11. Male aedeagus with broad, bilobed apex; parameres not connected, recurved 

apices long and scimitar shaped; female abdomen pruinose gray above, with 
narrow, shining brown bands broadening at margins of segments. 

10. multilineata (Lutz) 

Male aedeagus cleft one-third way to base, with a pair of long, slender, apical 

points; parameres broadly connected at bases, recurved apices with broadly 

rounded lobes 11. guianae, new species 

12. Mesonotum pruinose gray with dark-brown spots or irregular patches; scutel- 

lum yellow in middle, with about 8 marginal hairs; wing bands distinct to 

hind margin, usually strongly interconnected 12. nebulosa (Coquillett) 

Mesonotum dark brown with irregular, pruinose, gray spots; scutellum usually 
entirely black, with 4 marginal hairs; wing bands faint behind vein Mj, 
usually rather well separated 13. johannseni, new species 

Subgenus Schizohelea Kieffer, 1917 

Schizohelea Kieffer, Ann. Mus Nat. Hungarici, vol. 15, p. 295, 1917, vol. 16, p. 57, 
1918, and vol. 17, p. 89, 1919.— Goetghebuer, Mem. Mus. Hist. Nat. 
Belgique, vol. 8, p. 65, 1920. — Edwards, Trans. Ent. Soc. London, vol. 74, 
p. 411, 1926.— Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 782, 1943. 
(Genotype, Cerafopogon copiosus Winnertz; monobasic.) 

Allohelea Kieffer, Ann. Mus. Nat. Hungarici, vol. 15, p. 364, 1917. (Genotype, 
Sphaeromyas pulchripennis Kieffer; original designation.) 



138 PROCEEDINGS OF THE NATIONAL MUSEUM ^°^- ^°» 

Following Goetghebuer (1934), I regard Schizohelea as a subgenus 
for leucopeza Meigen. 

Diagnosis. — This species differs rather markedly from the other 
species of Monohelea in having the second branch of the media broadly 
interrupted at the base, basitarsus without apical spine, and male 
genitalia with emarginate sternite and arched aedeagus. 

1. Monohelea (Schizohelea) leucopeza (Meigen), 1804 

Figure 18, m 

Ceratopogon leucopeza Meigen, Klassifikation und Beschreibung . . . Insekten, 

vol. 1, p. 29, 1804. 
Ceratolophus leucopeza, Kieffer, Genera insectorum, fasc. 42, p. 60, 1906. 
Schizohelea leucopeza, Edwards, Trans. Ent. Soc. London, vol. 74, p. 411, 1926. — 

Seguy, Faune de France, pt. 8, p. 70, 1937. 
Monohelea (Schizohelea) leucopeza, Goetghebuer, in Lindner, Die Fliegen der 

Palaearktischen Region, Lief. 78, p. 53, 1934. 
Ceratopogon albiiarsis Wiedemann, Zool. Mag., vol. 1, p. 67, 1817. 
Ceratopogon copiosus Winnertz, Linn. Ert., vol. 6, p. 56, 1852. 
Ceratolophus copiosus, Kieflfer, Genera insectorum, fasc. 42, p. 60, 1906. 
Schizohelea copiosa, Kieflfer, Ann. Mus. Nat. Hungarici, vol. 15, p. 295, 1917; idem, 

vol. 16, p. 57, 1918; idem, vol. 17, p. 89, 1919.— Goetghebuer, Mem. Mus. 

Hist. Nat. Belgique, vol. 8, p. 66, 1920. — Kieffer, Faune de France, pt. 11, 

p. 116, 1925. 
Ceratopogon politus Coquillett, Proc. U. S. Nat. Mus., vol. 23, p. 606, 1901 ( 9 , 

Massachusetts) . 
Ceratolophus politus, Kieflfer, Genera insectorum, fasc. 42, p. 61, 1906. 
Johannseniella polita, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 

227, 1914. 
Johannsenomyia polita, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 

335, 1915 (New York). 
Sphaeromias polita, Kieflfer, Ann. Mus. Nat. Hungarici, vol. 15, p. 364, 1917. 
Allohelea polita, Kieflfer, Ann. Mus. Nat. Hungarici, vol. 15, p. 364, 1917. 
Schizohelea polita, Johannsen, Ann. Ent. Soc. Amer.. vol. 36, p. 782, 1943 

(? ^leucopeza Meigen). 
Bezzia stecki Kieflfer, Broteria, Ser, Zool., vol. 13, p. 65, 1915. 

Description. — Female: Length 1.5 mm., wing 1.2 mm. by 0.5 mm. 
Entirely shining black, only the tarsi, wings, and halteres white. 
Antennae quite short, the distal segments scarcely elongated. 
Mesonotum with scattered, erect, long, black hairs; scutellum with 
six long marginal hairs. Second radial cell half again as long as the 
first, M2 broadly interrupted at base. Spermathecae two, subequal, 
rather small and subspherical, the ducts sclerotized a short distance. 

Male (based on specimen from Suffolk, England): Ninth sternite 
about three times as broad as long, with a shallow, round emargination 
in middle of caudal margin, not spiculate; ninth tergite narrow, sur- 
passing basistyles, distal half with margins subparallel, apex truncate 
with setose apicolateral corners, the inner surface coarsely spiculate. 
Basistyles broad at base, each with a distinct lobe bearing three setose 
tubercles on inner side; abruptly narrowed on distal half; dististyles 



HELEID MIDGES, GENUS MONOHELEA — WIRTH 139 

Table 1. — Proportions of segments of hind legs of female Monohelea species 



Species 


Cx 


Tr 


F 


Ti 


Ti 


T2 


T3 


T4 


T5 


1. leucopeza 

2. hieroglyphica 

3. maculipennis 

4. lanei 


25 
30 
20 
25 

20 
24 
30 

28 
28 
25 


10 
10 
10 
10 
10 
8 
10 
12 
12 
10 


65 
80 
55 
65 
60 
72 
80 
88 
80 
70 


65 
72 
50 
60 
55 
68 
70 
80 
70 
65 


30 
40 
24 
35 
30 
33 
50 
50 
24 
25 


12 
16 
11 
17 
15 
16 
20 
20 
12 
12 


10 

9 

8 

10 

10 

9 

12 

12 

6 

8 


8 
8 
6 
8 
7 
7 
8 
8 
8 
10 


12 
8 
4 
8 


5. macfiei 


7 


7. ornata 

9. stonei 


7 
10 


10. multilineata 

12. nebulosa 

13. johannseni.. 


8 

8 

10 







nearly as long as basistyles, rather strongly curved. Aedeagus in 
form of a slender, rounded arch nearly as long as broad at base, with 
a pair of short, pointed, submedian plates projecting dorsocaudad 
at apex. Parameres with stout, lateral apodemes at bases; stems 
stout and gently curved, contiguous at midlength, their apices pointed 
and abruptly bent ventrocephalad on about distal fourth. 

Type. — Presumably in the Meigen collection in the Museum 
National d'Histoire Naturelle in Paris. 

Material examined. — Massachusetts: N. Amherst, June 1940, 4 99. 

New York: Mecklenburg, June 2, 1940, A. Stone, 10 99; North 
Ridgeway, June 22, 1940, S. C. Mendall, 1 9. 

Michigan: St. Joseph, May 30, 1938, C. W. Sabrosky, 6 99. 

England: Corriegills, Arran, June 2-4, 1919, F. W. Edwards, 2 99; 
Mildenhall, Suffolk, May 22, 1909, Yerbiu-y, 1 &. 

Remarks. — This species is foimd in Europe, West Africa, and 
northeastern North America. 

Subgenus Monohelea Kieffer, 1917 

Monohelea Kieffer, Ann. Mus. Nat. Hungarici, vol. 15, p. 295, 1917. — Goetghebuer, 
Mem. Mus. Hist. Nat. Belgique, vol. 8, p. 63, 1920. — Ingram and Macfie, 
Ann. Trop. Med. Parasit., vol. 15, p. 344, 1921. — Edwards, Trans. Ent. Soc. 
London, vol. 74, p. 410, 1926.— Tokunaga, Tenthredo, vol. 3, p. 156, 1940.— 
Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943 — Lane, Arq. Fac. Hig. 
Saude Univ. Sao Paulo, vol. 1, p. 225, 1948. — Lee, Proc. Linn. Soc. New 
South Wales, vol. 72, p. 350, 1948. (Genotype, Monohelea hieroglyphica 
Kieffer; original designation.) 
Diagnosis. — Wings with prominent markings; body with pruinose 

pattern; ninth stemite of male spiculate and transverse or lobed, not 

emarginate. 

hieroglyph ica — group 

Diagnosis. — Hind legs conspicuously banded, not markedly swollen; 
hind tarsi slender; wings with extensive irregular markings; male 
aedeagus broad, with median, anterior notch and triangular, lateral 
sclerites with converging, sharp-pointed apices. 



140 PROCEEDINGS OF THE NATIONAL MUSEUM ^^^- "^ 

2. Monohelea {Monohelea) hieroglyphica Kieffer, 1917 

Figure 19, g 

Monohelea hieroglyphica KieflFer, Ann. Mus. Nat. Hungarici, vol. 15, p. 312, 1917 

(cf , 9 , Paraguay). 
Monohelea hyeroglyphica, Lane, Arq. Fac. Hig. Saude Pub. Univ. Sao Paulo, vol. 

1, p. 225, 1948 (Brazil). 

Discussion. — This is the genotype of Monohelea. Macfie's (1937, 
1940a) records of this species from Trinidad and British Guiana must, 
I beUeve, be referred to macfiei, new species, described (see. p. 143), 
from Louisiana. 

Through the kindness of John Lane I have examined a female from 
km. 47, estrada Rio-Sao Paulo, Rio de Janeiro, Brazil, collected in 
February 1945 by P. Wygodzinsky. This is from the same locality 
as the male described and figured by Lane. Externally this species is 
almost inseparable from M. (M.) lanei, new species (see p. 142), from 
Florida, but the male genitalia are very close to those of M. (M.) 
maculipennis. A sketch of the male parameres (after Lane) is included 
here for comparison with those of maculipennis. 

Types. — In the Musei Nationalis Hungarici in Budapest, cf, 9, 
Paraguay. 

3. Monohelea {Monohelea) maculipennis (Coquillett), 1905 

Figures 18, a, i; 19, / 

Ceratopogon maculipennis Coquillett, Journ. New York Ent. Soc, vol. 13, p. 64, 
1905 (9 , Florida). — Kieffer, Genera insectorum, fasc. 42, p. 51, 1906. 

Monohelea maculipennis, KieflFer, Ann. Mus. Nat. Hungarici, vol. 15, p. 312, 1917. — 
Johannsen, Ann. Ent. Soc. Amer. vol. 36, p. 781, 1943. 

Diagnostic characters. — A yellowish gray, pruinose species with 
faint, brown, mesonotal dots, pale j^ellowish scutellum; hind legs with 
narrow, dark rings and mngs with extensive, irregular, grayish macula- 
tions bordered with whitish pruinosity. 

Description. — Female: Length 1 mm., wing 1 mm. by 0.4 mm. 
Head pale yellowish, antennae and palpi bro\^ii ; proportions of flagellar 
segments of antennae 12:10:10:12:12:12:12:12:16:16:18:18:24. 
Palpal segments in proportion of 5 :7 : 10 :5 : 10. 

Mesonotum grajdsh pruinose, with many scattered, small, brown 
dots at the bases of the dark mesonotal haks; humeri and sides exten- 
sively yellow. Scutellum pruinose, yello-wish white, with four marginal 
hairs, the middle pair quite close together. Postscutellum and pleura 
pruinose brown, the latter with several indistinct, transverse, darker 
lines. Legs yellowish, coxae and trochanters brown, upper fourth of 
midcoxa and hind coxa yellowish ; fore femur and midf emur and tibiae 
mibanded ; hind femur (figure 18, i) brown at extreme base, two narrow 
oblique dark rings in middle and a dark, preapical, ventral spot; hind 
tibia mth narrow sub-basal, median, and apical rings; tarsi narrowly 



HELEID MIDGES, GENUS MONOHELEA — WIRTH 141 

dark at apices of segments. Proportions of segments of hind legs as in 
table 1. 

Wing whitish hyaline, with irregular maciilations, grayish by 
transmitted light, yellowish bro^vn with narrow, whitish pruinose 
borders by reflected light. Markings composed of a broad, irregular 
band across wing at level of first radial cell, filling basal half of medio- 
cubital fork, with a prominent, omega-shaped spur in base of cell Rg; 
a small spot before middle of second radial cell; a more or less quadrate 
to X-shaped mark across cell R5 at apex of second radial cell ; a sigmoid, 
subapical mark across cells Mi and M2, often connected by very narrow 
lines in these cells to mesal band ; small, irregular spots across wing near 
base and a small spot past middle of basal cell. First radial cell about 
half as long as second, a very few macrotrichiae at apices of cells R5 
and Ml, Halteres dull white, a black dot on anterior side. 

Abdomen dull whitish, sides with irregular, dark patches. Sper- 
mathecae two, very unequal, subspherical, each with short sclerotized 
duct. 

Male: Ninth sternite about twice as broad as long, spiculate, 
posterior margin transverse, abutting against base of aedeagus, with 
four long hau*s arising from tubercles in a curved row; ninth 
tergite greatly constricted, with sides subparaUel on distal half, apex 
truncate, apicolateral processes short. Basistyles narrowed on distal 
halves ; dististyles curved to slender, pointed apices. Anterior margin of 
aedeagus broad, slightly concave, with a small median notch; aedeagus 
bearing a pau- of triangular, submedian sclerites, the bladelike apices 
of which converge before level of the irregular bilobed apices of an 
accessory pair of dorsal sclerites. Parameres with flaring, winglike, 
bilobed, basal apodemes, more or less connected at midlength by a 
pair of stout mesal lobes, apices each with a straight, long, slender 
stem bearing a dorsolateral lobe about half as long and of same thick- 
ness, just beyond level of mesal bridge. 

Type.—VSlS^M 8366, 9, Florida, Jacksonville. 

Material examined. — Florida: Crystal River, Citrus County, July 
17, 1950, Hudson, 4 99; Everglades City, Collier County, Feb. 7, 
1950, Davidson, 1 cf^, 9 99; Fort Myers, Lee County, Feb. 2, 1949, 
Brechtel, 1 cf ; Islamorada, Monroe County, June 7, 1949, Smith, 1 
cT, 2 99; Jacksonville, date not given, Slosson, 1 9 (type). 

Mexico: Ciudad Monte, Tamaulipas, Nov. 22, 1943, Brookman, 1 
&,S 99. 

Guatemala: Rio Dulce, Mar. 21, 1906, Schwarz and Barber, 1 9. 

Panama: Rio Trinidad, June 9, 1902, Busck, 1 9. 

Remarks. — Formerly all the North American Monohelea with the 
type of wing pattern characteristic of hieroglyphica were called 
maculipennis. Thus in my (1952) paper on California Heleidae, I 



142 PROCEEDINGS OF THE NATIONAL MUSEUM ^°^- "^ 

erroneously included a California record of maculipennis and also 
cited several eastern records of this species in error. I have now 
studied the hieroglyphica group more intensively and believe I have 
correctly identified several component species on the basis of male 
genitalia supported by several rather difficult but constant external 
characters of the female. In addition to hieroglyphica and maculi- 
pennis there appear to be four or more undescribed species in this 
complex. Although my California specimen and that reported as 
Monohelea sp. by Johannsen (1943) do not fit any of those described 
below, probably falling closest to hieroglyphica in wing markings, a 
positive diagnosis of the species must await the collection of the male. 

4. Monohelea (Monohelea) lanei, new species 

Figures 18, b, j; 19, e 

Description. — Female: Length 1.3 mm., wing 1.1 mm. by 0.4 mm. 
As in M. (M.) maculipennis (Coquillett), but darker and slightly 
larger. Mesonotum light brown pruinose with faint brown punctures; 
scutellum brownish in middle. Markings on hind legs stronger; 
basal third of hind femur brown, a narrow, diagonal band just past 
middle, then a small, brown, ventral spot and a strong preapical band; 
basal and median bands on hind tibia broad and joined on ventral side, 
the distal band broad. Proportions of segments of hind leg as in 
table 1. 

Wing markings darker and more extensive than in maculipennis, 
the omega-shaped mark entirely closed behind by a broad extension 
of the mesal band, the X-shaped mark in cell R5 greatly constricted in 
middle and much broader behind. Macro trichiae sparse and con- 
fined to wing margin; first radial cell about two-thirds as long as 
second. 

Abdomen brown above, yellowish on sides in front and on three 
distal segments. Spermathecae two, large, very unequal and sub- 
spherical. 

Male: Ninth sternite with posterior margin convex in middle, 
spiculate; ninth tergite tapered to a pair of blunt, triangular, sub- 
median, caudal lobes. Basistyles narrow, nearly straight; dististyles 
slender, curved, with sharp, incurved points. Aedeagus with basal 
halves broadly separated by a deep notch in middle of anterior margin, 
lateral sclerites with broad, triangular bases, the slender, pointed, 
posterior portions directed caudomesad and meeting each other over 
tips of a pair of sharp-pointed, dorsal processes. Parameres a pair 
of irregular, sublateral sclerites joined by an indistinct transverse 
bridge at a third of the distance from bases; basal apodemes trilobed, 
directed laterocephalad to bases of basistyles; posterior portions with 
subparallel inner margins, tips each with apical, knoblike, sclerotized 
lobe with slightly longer, flattened, roundly flaring, ventral expansions. 



HELEID MIDGES, GENUS MONOHELEA WIRTH 143 

Types. —Uolotype, a", USNM 61091, Miami, Florida, Mar. 25, 
1944, Wirth. Allotype with same data as type except date, October 
1943. Paratypes: Florida: 2 d* d^, 59 9 , same, except date, October 
1943 to April 1944; 1 9, Welaka, June 17, 1946, Bellamy; 1 9, 
Everglades City, Feb. 7, 1950, Davidson. 

5. Monohelea (Monohelea) macfiei, new species 

Figures 18, c, h; 19, h 

Monohelea hieroglyphica, Macfie, not Kieffer (misicjentification), Ann. Mag. Nat. 
Hist., ser. 10, vol. 20, p. 18, 1937 (9 , Trinidad); Ent. Monthly Mag., vol. 
76; p. 30, 1940 (cf, British Guiana, genitalia figured). 

Description.- — Female: Length 1.2 mm., wing 1.4 mm. by 0.5 mm. 
As in M. (M.) maculipennis (Coquillett), but the mesonotum with a 
pronounced, broad, median, anterior band and a large area between 
wing bases in front of scutellum uniform brown, the scutellum with a 
more or less brownish median area. Abdomen suffused, grayish 
brown above. Legs as figured by Macfie (1937), brown bands on 
hind legs quite distinct; on hind femur a band on basal third, a diagonal 
band past middle, and a narrov/ band subapically ; on hind tibia, 
narrow basal, median, and apical rings; all tarsal segments with 
narrow apical bands. 

Wing as in figure 18, c, the dark areas in the first basal and anal cells 
quite extensive, the X-shaped mark in cell R5 large, with the caudo- 
distal arm more prolonged than shown by Macfie and the distal sig- 
moid mark in cells Mi and M2 connected by broad lines in these cells 
to the broad median crossband. All wing markings brown by re- 
flected light, gray by transmitted light; a few macro trichiae at apex 
of wing. 

Male: The resemblance of the Louisiana male to the male from 
British Guiana figured by Macfie (1940a) is remarkable. The 
slender, sinuate, bladelike parameres connected hy a narrow bridge 
at midlength are the most distinctive difi^erence. 

Types. — Holotype, cf ,USNM 61092, allotype, Kilbourne, Louisiana, 
May 10, 1947, Wirth. Paratypes: Louisiana: 2 9 9, with same data 
as type; 1 9, Baton Rouge, May 16, 1947, Wirth. 

6. Monohelea {Monohelea) texana, new species 

Figure 19, d 

Description. — Male: Wing 1.1 mm. long. Badly damaged, anten- 
nae, all of legs but fore femur, midfemm", and tibiae gone. Mesonotum 
appears discolored, all dark except large quadrate yellow patches on 
humeri; scutellum light yellow, a minute brown spot in middle of 
anterior surface. Forelegs and midlegs yellow, narrow basal, median, 
and subapical faint brown rings on fore femur; nan'ow basal and 

232992—53 2 



144 PROCEEDINGS OF THE NATIONAL MUSEUM ^°^- ^'^ 

broader median and apical brown rings on fore tibia; broad basal and 
narrow median brown rings on midfemur and faint median and 
broad, dark-brown apical rings on midtibia. Wings marked as in 
M. (M.) hieroglyphica Kieffer, the median band broad, the omega- 
shaped spur distinct and open below, the spot in cell Rs narrow with 
the right-hand arms of the X absent, the sinuate mark in cells 
Ml and Mj reduced to three discrete spots; macro trichiae reduced to 
a few at v/ing margin. Halteres yellow with flat ends a.nd a spot on 
anterior surfaces of knobs dark. 

Ninth sternite spiculate, posterior margin broadlj'^ convex in middle, 
with four long hairs arising from bases in an arched row; ninth tergite 
tapered, with a pair of prominent, apicolateral processes. Aedeagus 
with basal halves narrowly separated by a deep notch in middle of 
anterior margin, lateral sclerites with broad, triangular bases, the 
slender, pointed posterior apices meeting each other at about level 
of tip of ninth tergite; the dorsal accessory structure biiid a third 
the way to base of aedeagus, with a pair of very slender lobes with 
pointed apices curved ventrad and surpassing apices of lateral sclerites. 
Parameres joined together broadly near middle, each with well- 
developed basal and lateral arms, stems abruptly bent laterad two- 
thirds the way to apices, then abruptl}'' bent ventrad, with pointed 
apices directed mesocaudad. 

Type. — Holotype, cf , Cornell Univ. Type 2787, Limpia Canyon, 
Davis Mountains, Texas, July 7, 1917, Bradley. 

Remarks. — The male genitalia are most distinctive, the simple 
apices of the parameres allying M. (M.) texana with M. (M.) macfiei, 
new species. However, the external features, as nearly as can be 
ascertained from the badly damaged specimen, are closer to other 
species, the wing markings being nearly like those of M. (M.) hiero- 
glyphica Kieffer, while the front legs and midlegs are banded as in 
M. (M.) ornata, new species, and M. (M.) brasiliensis Lane. 

7. Monohelea {Monohelea) ornata, new species 

Figure 18, d 

Description. — Female: Length 1.2 mm., wing 1.2 mm. by 0.5 mm. 

Head pruinose brown, vertex yellowish; antennae broken, pedicel 
brown; palpi brown. Mesonotum pruinose gray, humeri yellowish; 
a broad, median, longitudinal band and broad patches above wings 
rich, velvety brown. Scutellum pruinose, grayish yellow, brown in 
mJddle; postscutellimi and plemm pruinose dark brown. Fore femur 
and midfemur dull yellow, with faint median brown bands; fore tibia 
and midtibia brown; hind femur dark brown on basal third, a broad, 
diagonal, brown ring past middle and a narrow preapical brown ring; 



HELEID MIDGES, GENUS MONOHELEA — WIRTH 145 

hind tibia with broad basal and apical rings, and a band just before 
middle, brown; tarsi dull yellowish. Proportions of segments of 
hind legs as in table 1; hind femur and tibia with rather long hairs; 
hind basitarsus with strong basal spine. Claws on forelegs long and 
equal, nearly as long as fom-th and fifth segments combined; midtarsi 
broken ; on hind leg a single long claw half again as long as fifth segment. 

Wing grayish hyaline, with very extensive gray maculations as in 
figure; maculations of hieroglyphica type, but much more extensive, 
with an extra distal, separate spot narrowly connected to the X-shaped 
mark in cell R5, and the subapical mark in cells Mi and M2 quite broad. 
First radial cell about half as long as second; macrotrichiae very ex- 
tensive, sparsely covering distal third of wing and including most of 
cell M 3 4. 4 and anal cell. Halteres not visible. Abdomen uniformly 
dark, pruinose brown, spermathecae not examined. 

Types.—Holotjpe, 9, USNM 61093, Santa Rosa Island, Escambia 
County, Florida, Oct. 10, 1949, Butler. 

Remarks.-— The wing, mesonotal, and leg markings are so distinctive 
that I do not hesitate to describe M. (M.) ornata from the single female. 

8. Monohelea {Monohelea) brasiliensis Lane, 1948 

Monohelea brasiliensis Lane, Arq. Fac. Hig. Saude Pub. Univ. Sao Paulo, vol. 1, 
p. 226, 1948 (9, Brazil). 

Description. — ^I have not seen this species, which is known only from 
the type female. Length 1.2 mm., wing 1.3 mm.; mesonotum chest- 
nut, yellowish on sides and in prescutellar depression; scutellum 
whitish. Legs yellowish, fore femur dark at base, mudfemur dark on 
basal half; fore tibia and mid tibia dark at apices; hind femur with two 
dark rings, one at apex of basal third and one in middle, hind tibia dark 
at base and apex and indistinctly so in middle. Hind basitarsus 1.8 
times as long as second segment. Wing with about 20 black dots in 
addition to irregular grayish markings; macrotrichiae numerous on 
distal third of wing; first radial cell slightly over half as long as 
second (from original description). 

Type. — In collection of University of Sao Paulo, Brazil, No. 6781, 
9, Brazil, Rio de Janeiro, Estrada Rio-Sao Paulo, km. 47. 

midnlineata — group 

Diagnosis. — Wings with two large anterior dark patches and other 
fainter irregular markings; scutellum dark in middle; legs dark with 
yellow laiee spots; hind legs slender, the tarsi long and unspined; 
ninth sternite of male with median, convex lobe and several long 
hairs, aedeagus long and narrow, parameres very long, with sharp 
apices. 



146 



PROCEEDINGS OF THE NATIONAL MUSEUM 




.LANEI 



k. JOHANNSENI 



W.LEUCOPEZA 



Figure 18. — Species of Monohelea: a-g, Wings: a, maculipennis (Coquillett); b, land, new 
species; c, macfiei, new species; d, ornata, new species; e, stonei, new species; /, nebulosa 
(Coquillett); g, johannseni, new species, h-k, Hind legs of females: h, macfiei, new species; 
i, maculipennis (Coquillett); ;', lanei, new species; k, johannseni, new species. /, Sperma- 
thecae, johannseni, new species, m, Male genitalia, leucopeza (Meigen). 



HELEID MIDGES, GENUS MONOHELEA — ^WIRTH 147 




STONEI 



J. MULTILINE ATA 



*. GUIANAE 



Figure 19. — Species of Monohelea, male genitalia; a, johannseni, new species; h, tessellata 
(Zetterstedt); c, nehulosa (Coquillett); d, texana, new species; e, lanei, new species; /, 
maculipewiis (Coquillett); g, hieroglyphica Kieffer; male parameres (after Lane); h, macfiei, 
new species; t, stonei, new species; ;, multilineata (Lutz); k, guianae, new species. 



148 PROCEEDINGS OF THE NATIONAL MUSEUM ^°^- "^ 

9. Monohelea {Monohelea) stonei, new species 

Figures 18, e; 19, i 

Diagnostic characters. — A yellowish gray, pruinose species with 
maculate wings; scutellum yellow with brown center; legs dark, 
abdomen dull gray pruinose above. 

Description. — Female: Length 2.0 mm., wing 1.5 mm. by 0.7 mm. 
Head yellowish, vertex pruinose, pale gray; antenna with pedicel and 
bases of flagellar segments yellow, apices of short segments and all of 
long distal segments and palpi brown; proportions of flagellar seg- 
ments 15:10:10:10:10:10:12:12:18:18:18:18:24. Palpal segments 
in proportion of 5 :8 : 15 : 10 : 15, third segment scarcely swollen, with a 
rather large sensory pit at midlength. 

Mesonotum yellowish gray pruinose, with tiny brown dots at the 
bases of the brownish mesonotal hairs. Scutellum yellow, brown in 
middle, with four moderate marginal hairs and a few fine setae. 
Postscutellum and pleura pruinose dark brown. Coxae brown; 
trochanters, femora, and tibiae light brown on forelegs and midlegs, 
black on hind legs; all knees prominently yellowish, tarsi yellow. 
Hind femur and hind tibia moderately thickened; basitarsus with a 
slender spine at base, distal tarsal segments on hind legs each with 
a fine apical spine; a single long claw on each hind leg. Proportions 
of segments of hind leg as in table 1 . 

Wing grayish hyaline, with two prominent, black spots and fainter 
posterior infumation as figured; fu-st radial cell about half as long as 
second. Halteres infuscated. 

Abdomen dark brown, entire dorsum pruinose gray except for a 
pair of small lateral spots and a pair of submedian sensory dots on 
each segment, polished brown. Spermathecae two, subequal, ovoid, 
each with a very short sclerotized duct. 

Male: Similar to the female, with the usual sexual differences; 
antennal plume golden at base, dark brown toward apex. Ninth 
sternite three times as broad as long, with a narrow, median, poste- 
rior lobe bearing two long hau-s extending into the concave base of the 
aedeagus; sternite spiculate on posterior half; ninth tergite as long as 
basistyles, tapered to the truncated apex, bearing a pair of short, 
rounded, apicolateral processes. Basistyles about twice as long as 
broad, simple, very convex on outer sides; dististyles about 0.7 as long 
as basistyles, nearly straight and stout to apices. Aedeagus with 
base as broad as the lobe of ninth sternite, a slight sub-basal swelling, 
a second swelling at midlength, with distal half cleft into a pair of 
attenuated, submedian, pointed blades attaining 0.8 length of basi- 
styles. Parameres consisting of a pau' of very irregular processes. 



HELEID MIDGES, GENUS MONOHELEA — ^WIRTH 149 

bases of each with a large anterior and lateral apodeme, stem rather 
stout and crooked, distal portion abruptly bent ventrocephalad, then 
greatly attenuated and bent ventrocaudad to a sharp point. 

Types. — Holotype, 9, USNM 61094, Baton Rouge, Louisiana, May 
16, 1947, Wirth. Allotype with same data as type except date. 
May 3, 1947. Paratypes: Louisiana: ecfcf , 3 99, with same data as 
t3npe except date, May 6-20, 1947. Mississippi: 4 cT" d", 10 99, Horn 
Island, June and July 1944, E. A. Richmond. Florida: 11 99, Gray- 
ton Beach, Walton County, May, Sept., 1949, Butler; 1 9, Santa 
Rosa, Walton County, July 19, 1949, Peterson; 8 cf cT, 21 99, In- 
nerarity Point, Escambia County, May, 1950, Rathert; 8 99, Santa 
Rosa Island, Escambia County, July 7, 1949, Butler; 5 99, Panama 
City Beach, Bay County, May 6, 1949, McElvey; 5 99, Crystal River, 
Citrus County, July 17, 1950, Hudson; 3 99, Fort Myers, Lee County, 
Oct. 8, 1948, Brechtel; 3 99, Everglades City, May, Oct., 1948, 1950, 
Huntoon; 1 cf , 1 9, Big Pine Key, Monroe County, Apr. 1, 1950, 
Sermon. Georgia: 2 cf cf, 4 99, Thomasville, May 15-30, 1949, 
Palmer. Tennessee: 1 9, Nashville, June 18, 1937, Adams. Iowa: 
1 9, Sioux City, July 15, 1950, Laffoon. Virginia: 3 d" cf^, 6 99, Falls 
Church, July 8, 29, 1950, Wirth. Maryland: 4 99, Leeds, Dorchester 
County, July 10, 1907, Barber. New Jersey: 1 9, Newport, July 1, 
1937 (light trap). Costa Rica: 1 cf , Higuito, San Mateo, no date, 
Schild. Panamd: 1 9, Fort Kobbe, Canal Zone, Aug. 21, 1950, 
Carpenter. 

10. Monohelea (Monohelea) multilineata (Liitz), 1914 

Figure 19, j 

Palpomyia multilineata Lutz, Mem. Inst. Oswaldo Cruz, vol. 6, p. 93, 1914 (Brazil). 
Monohelea multilineata, Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943. — 
Lane, Rev. Ent., vol. 16, p. 368, 1945. 

Lane (1945) has very adequately redescribed this Brazilian species. 
It resembles M. (M.) stonei, new species, in general coloration and 
structure, the mesonotum yellowish with brown dots, the scutellum 
yellow with median brown spot and four marginal bristles, forelegs 
and midlegs yellowish, the hind femur and tibia black and scarcely 
swoUen and hind tarsi unspined. The wings, however, in addition 
to the two anterior black spots, have smaller, rather strong spots at 
the wing margin m cells R5 and Mi, and across the apex of cell M2. 
The polished, brown, lateral spots on the abdomen, which are small 
in stonei, are larger and narrowly connected across the anterior margin 
of each tergite in multilineata. 

The male genitalia of multilineata are of the same type as those of 
stonei, but there are four rather than two bristles at the base of the 



150 PROCEEDINGS OF THE NATIONAL MUSEUM ^®^- ^<" 

lobe of the ninth sternite, the aedeagus is broader at the apex and 
only slightly notched rather than deeply cleft, and the parameres are 
slender and nearly straight with the apices abruptly bent and scim- 
itar-shaped. 

Material examined. — Brazil : Km. 47, Estrada Rio-Sao Paulo, Rio de 
Janeiro, Brazil, February 1945, Wygodzinsky, 2 cfcf , 2 99. 

Remarks. — Monohelea nigeriae Ingram and Maciie, 1922, known 
from the female from West Africa, is closely related to M. (M.) stonei 
and to M. (M.) muUilineata, but according to the original description, 
the tarsi bear ventral spines, as in the tessellata group, and the sperma- 
thecae are imequal in size. 

11. Monohelea (Monohelea) guianaef new species 

Figure 19, A; 

Monohelea muUilineata, Macfie, not Lutz (misidentification) , Proc. Ent. Soc. 
London, Ser. B, vol. 9, p. 187, 1940 (c?, British Guiana). 

Macfie (1940) has given as satisfactory a description of this insect 
as was possible from the single male specimen available. This speci- 
men was borrowed from the British Museum for study and externally 
cannot be separated from either M. (M.) muUilineata (Lutz) or 
M. (M.) stonei, new species. However the male genitalia are quite 
distinct and may be characterized as follows: Ninth sternite very 
short, with a low median lobe on caudal margin fitting against base of 
aedeagus, spiculose, with four long hairs arising from tubercles in a 
line at base of median lobe; nmth sternite very narrow, tapered to 
tip, with a pair of short, nipplelike apieolateral processes. Basistyles 
about twice as long as broad; dististyles nearly straight, with blimt, 
scarcely narrowed apices. Aedeagus very narrow, about 2.5 times as 
long as broad at base, the anterior arch very low, apex cleft about a 
third way to base, with a pair of very slender, submedian processes, 
from the bases of which a pair of slender, lateral arms curve around 
stems of parameres. Parameres broadly joined together at bases, 
each with a short anterior and lateral arm, stems slender, apex of each 
thickened and abruptly bent laterocephalad in a broadly rounded 
lobe about half as long as dististyle. 

Types. — Holotype, cf, British Museum (Natural History) collec- 
tion, Mazaruni, British Guiana, Aug. 21, 1937, second growth (low 
forest), Richards and Smart. 

tessellata — group 

Wings with three large dark anterior spots and irregular posterior 
infuscated areas; scutellum often with middle yellow; hind legs swollen 
and black, with yellow knees, the tarsi short and with strong ventral 



HELEID MIDGES, GENUS MONOHELEA — WIRTH 151 

spines; claw of hind leg of male long and single as in the female; male 
aedeagus with slender, pointed, posterior sclerite borne in a notch in 
the quadrate basal sclerite, parameres short, curved, and pointed. 

12. Monohelea {Monohelea) nebulosa (Coquillett), 1901 

Figures 18,/; 19, c 

Ceratopogon nebulosus Coquillett, Proc. U. S. Nat. Mus., vol. 23, p. 606, 1901 

(cf, New Jersey). 
Ceratolophus nebulosus, Kieffer, Genera insectorum, fasc. 42, p. 60, 1906. 
Johannseniella nebulosa, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, 

p. 226, 1914. 
Hartornyia nebulosa, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 340, 

1915 (Indiana). 
Monohelea nebulosa, KieflFer, Ann. Mus. Nat. Hungarici, vol. 15, p. 312, 1917. — 

Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943 (Massachusetts, 

Idaho) . 

Description. — Female: Length 2.1 mm., wing 1.7 mm. by 0.6 mm. 
Head dull, dark brown, with a few pruinose gray spots on vertex; 
antennae light brown, distal segments not darker. Palpi light brown. 

Mesonotum pruinose gray, with scattered, small, dark-brown dots, 
each mesonotal hair arising from a dot, these dots more or less con- 
fluent, especially on posterior portion of mesonotum. Scutellum 
brown, middle third yellow, with about eight marginal bristles. 
Postscutellum and plem-a dark brown. Legs shining dark brown, 
knees and tarsi yellowish. Posterior femora and tibiae greatly 
thickened; hind basitarsus with basal and distal spine; second segment 
with two distal spines, third segment with one distal spine; claws 
minute and equal on forelegs and midlegs, outer claw very long, 
four times the length of inner, on hind legs. Proportions of segments 
of hind leg as in table 1. 

Wing grayish hyaline, a quadrate black patch across first basal cell 
at half its length; two broad, irregular, more or less interconnected, 
transverse bands across wing, the first at first radial cell, the second at 
apex of second ; cell Rs thus with a dark, a light, a dark and a light band, 
all of subequal breadth. First radial cell 0.4 times as long as second. 
Halteres pale yellow. 

Abdomen uniform, pruinose, dark brown; anal segment yellow. 

Male: Ninth sternite transverse, without caudal lobe, posterior 
portion spiculate; ninth tergite very narrow, with sides subparallel on 
distal half, apicolateral processes short and rounded. Basistyles very 
stout and convex; dististyles slender and curved, 0.6 as long as basi- 
styles. Aedeagus with two sections, basal part about as broad as long, 
anterior margin contiguous with sternite, posterior margin deeply 
emarginate, the slender, rodlike, distal sclerite with notched apex fit- 



152 PROCEEDINGS OF THE NATIONAL MUSEUM ^°^- ^°^ 

ting into the emargination. Parameres connected by a transverse 
bridge near bases, with slender anterior and lateral apodemes; each 
paramere with posterior portion narrowed and curved ventrolaterad 
to a bladelike apex, a small sub-basal tooth on ventral side. 

Material examined.— New York: Farmingdale, July 9, 1938, H. and 
M. Townes, 1 cf ; Millwood, June 21, 1936, H. Townes, 1 9. 

New Jersey: Atsion, June 17, 1949, H. Townes, 1 9; Medford Lakes, 
June 17, 1939, H. Townes, 1 cf; Riverside, June 18, 1939, H. Townes, 
1 9; Riverton, June 19, Johnson, 1 cf (type). 

Georgia: Thomasville, May 15-30, Pahner, 1 9. 

Arkansas: Pike County, June 12, 1938, Turner, 1 9. 

Remarks. — This species is very similar to the Palearctic species M. 
(M.) tessellata (Zetterstedt), 1850, which, however, has on the scutel- 
lum four marginal bristles, the ends of which are yellowish, as is the 
median area. The wing markings of tessellata are much more nearly as 
in M. (M.) johannseni, new species. 

A male of tessellata from Norfolk, Hatfield, England, was examined 
through the kindness of Paul Freeman of the British Museum (Natural 
History) . The specimen is unsatisfactory for a description of external 
characters, as it was glued to a card when wet. It is noted, however, 
that both the first and second segments of the hind tarsi bear a distal 
pair of spines. The aedeagus of the genitalia consists of a heavily 
sclerotized basal plate about three times as broad as long, bearing a 
small, crescentic, median sclerite on the ventroposterior margin and 
bearing dorsoposteriorly a large, triangular structure with sharp- 
pointed, median apex, and widely flaring, lateral arms articulating with 
inner margins of basistyles. Parameres with slender anterior arms 
connected by a broad, median bridge, the posterior portions simple, 
the basal halves stout, the distal halves very slender and curved ventro- 
cephalad. 

According to Tokunaga's (1940) description, his record of tessellata 
from Japan probably refers to an as yet unnamed species. This Jap- 
anese species has the scutellum yellow only in the middle, as in nebulosa, 
which it also closely resembles in wing markings; but the mesonotura 
has distinct yellow spots, the antennae have the distal segments and 
the apices of the basal segments brownish, the legs are yellowish brown 
with dark brown markings, and the dorsum of the abdomen is pale, 
yellowish brown. 

The West African species M. (M.) litoraurea Ingram and Macfie, 
1921, (female) and M. (M.) mimas de Meillon, 1939, (male) are very 
similar to M. (M.) nebulosa (Coquillett) . The genitalia of mimas are 
the same as those of nebulosa except that the dististyles are abruptly 
bent near the base. 



HELEID MIDGES, GENUS MONOHELEA — WIRTH 153 

13. Monohelea {Monohelea) johannseni, new species 

Figures 18, g, k, I; 19, a 

Monohelea tessellata, Johannsen, not Zetterstedt (misidentification) , Ann. Ent. 
Soc. Amer., vol. 36, p. 781, 1943 (Alabama). 

Diagnostic characters. — A dark brown species with large pruinose 
gray mesonotal markings, black scutellum and legs, and maculate 

WIQgS. 

Description. — Female: Length 1.5 mm., wing 1.2 mm. by 0.5 mm. 
Closely resembling M. (M.) nebulosa (Coquillett) but smaller, the 
mesonotum dark brown with large, irregular, pruinose gray areas, the 
scutellum entirely blackish and with four long and several shorter 
marginal hairs. Proportions of hind leg of female as in table 1, hmd 
tarsi with one basal and two distal spines on fu'st segment, two spines 
at tip of second and occassionally two spines at apices of third and 
fourth segments; inner claw on hind leg of female about a thu'd as 
long as outer. Wing as in nebulosa, but the markings behind vein 
iVli much fainter than those on anterior part of wing. Spermathecae 
two, subequal, oval, each with very short, sclerotized duct. 

Male genitalia as in nebulosa, but the parameres simple, without 
the sub-basal ventral tooth, and distinctly and abruptly bent mesad 
at distal fifth. 

Types.— Roloiype, 9, USNM 61095, Falls Church, Vkginia, July 8, 
1950, Wirth. Allotype, Innerarity Point, Escambia County, Florida, 
Apr. 29, 1949, Kathert. Paratypes: Virginia: 2 99, Mountain Lake, 
July 15, 1938, July 21, 1940, L. and M. Mihie. Florida: 4 cf cf , 5 99, 
same data as allotype except date April and May 1949; 1 9, Santa 
Rosa, Walton County, May 5, 1950, Peterson. Alabama: 1 9, La- 
Place, June 9, 1917, Bradley. Michigan: 1 9, Cheboygan County, 
July 17, 1942, Sabrosky. Iowa: 1 9, Pikes Peak State Park, Clayton 
County, July 4, 1949, Laffoon. 



References 

Floch, H,, and Abonnenc, E. 

1942. Ceratopogonides divers de la Guyane Francaise III, Inst. Pasteur 

Guyane Terr. L'Inini Publ. 55, 6 pp., 2 figs. 

GOETGHEBUER, M. 

1934. Heleidae (Ceratopogonidae), In Lindner, Die Fliegen der Palaearkti- 
schen Region, Lief. 78, pp. 49-94. 

JOHANNSEN, O. A. 

1943. A generic synopsis of the Ceratopogonidae (Heleidae) of the Anaericas, 

a bibliography, and a list of the North American species. Ann. 
Ent. Soc. Amer., vol. 36; pp. 763-791. 

KlEFFER, J. J. 

1917. Chironomides d'Amerique conserves au Musee National Hongrois de 
Budapest. Ann. Mus. Nat. Hungarici, vol. 15, pp. 292-364, 43 figs. 
Lane, J. 

1945. Redescrigao de Ceratopogonideos Neotr6picos (Diptera: Ceratopogo- 
nidae). Rev. Ent., vol. 16, pp. 357-372, 19 figs. 
Lee, D. J. 

1948. Australasian Ceratopogonidae (Diptera, Nematocera). Part 4. The 
Stilobezzia group of genera. Proc. Linn. Soc. New South Wales, 
vol. 72, pp. 345-356, 1 pi., 23 figs. 
Macfie, J. W. S. 

1937. Ceratopogonidae from Trinidad. Ann. Mag. Nat. Hist., ser. 10, vol. 

20, pp. 1-18, 6 figs. 
1940a. A report on a collection of Ceratopogonidae (Diptera) from British 

Guiana. Ent. Monthly Mag., vol. 76, pp. 23-32, 4 figs. 
1940b. Ceratopogonidae (Diptera) from British Guiana and Trinidad. 
Proc. Ent. Soc. London, ser. B, vol. 9, pp. 179-195, 4 figs. 
Tokunaga, M. 

1940. Chironomoidea from Japan (Diptera), XII. New or little-known 
Ceratopogonidae and Chironomidae. Philippine Journ. Sci., vol. 
72, pp. 255-311, 4 pi. 
WiRTH, W. W. 

1952. The Heleidae of California. Univ. California Publ. Ent., vol. 9, pp. 

95-266, 33 figs. 
154 



U S. GOVERNMENT PRINTING OFFICE: I9S3 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



issued sM^^vvL sj'l^i ^y 'A' 




SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 



Vol.103 Washington : 1953 No. 3321 



A REVIEW OF THE BEETLE FAMILY CEPHALOIDAE 



By Ross H. Arnett, Jr.^ 



The family Cephaloidae consists of only one genus, Cephaloon 
Newman, 1838, with eight known species that are locally rather com- 
mon but are known only from the Eastern United States, Western 
United States, Western Canada, Japan, and Amur. All of the species 
are closely related and quite variable in color. Useful separation 
characters are to be found in the antennae and various male structures. 
As yet Httle is known concerning the habits and life histories of mem- 
bers of this group. All known species of this family are represented 
in the collections of the U. S. National Museum. 

After study of some material from the Ussuri River Valley recently 
sent to me for determination, it seems desirable to reconsider the 
taxonomy, affinities, and distribution of this group. The ranking of 
this small assemblage of species as a family on equal ground with the 
rest of the families in the order is still somewhat doubtful in my mind. 
I believe the placing of the Cephaloidae as a satellite of the great 
family Tenebrionidae is firmly established, but my studies in the 
Tenebrionoidea have not as yet revealed, on the basis of our present 
ranking and evaluation of what defines a family, any group with which 
the genus Cephaloon can be incorporated; hence, I retain it as a family. 

I wish to thank Mr. Hugh B. Leech and Dr. E. C. Van Dyke, both 
of the California Academy of Sciences, and Dr. Hans Klapperich of 
Bonn, Germany, for the loan of several specimens used in this study. 

' Bureau of Entomology and Plant Quarantine: U. S. Department of Agriculture. 

232728—63 155 



156 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

Family Cephaloidae LeConte 

Cephaloidae LeConte, Smithsonian Misc. Coll., vol. 3, art. 3, Classification of the 
Coleoptera of North America, pt. 1, p. 259, 1862. 

LeConte originally proposed this family for the North American 
Cephaloon lepturides Newman, but also referred to two species, al- 
though not by name, which had been described by Motschulsky from 
the Amur River Valley. Several genera have since been erected for 
species in this group, and then synonymized, until in the present 
study the family contains only the original genus Cephaloon Newman. 

Family diagnosis. — Size 8-20 mm. ; head elongate, diamond-shaped, 
deflexed; antennae 11-segmented, filiform, with apical segments some- 
what enlarged, inserted between the eyes and bases of the mandibles; 
mandibles elongate, acute at apex, never bifid or otherwise modified; 
eyes reniform, not prominent; maxillary palpus 4-segmented, first 
segment small, obscure, apical segment triangular. Pronotum with- 
out lateral margins, smooth, always abruptly narrowed anteriorly 
from the middle. Legs slender; front coxal cavities open behind; 
front and middle coxae prominent, conical; apical spurs of all tibiae 
large, two on each tibia; tarsi 5-5-4, the segments all simple, not lobed 
or tomentose beneath; claws pectinate, with a subequal membranous 
lobe beneath each claw. Elytra with vague costae, minutely punctate, 
never striate. Abdomen with 5 visible sternites in the female, 7 in 
the male (sternites 2 +3 to 9 visible in the male). Body covered with 
very fine pubescence. 

Male genitalia: Apical abdominal segments (7-9) of the male 
considerably modified. Segment 7 with sternite and tergite laterally 
fused, forming a globular shaped segment. Segment 8 with sternite 
and tergite laterally fused, forming two laterally triangular pieces. 
Segment 9 closely fitted into segment 8, the tergite and sternite both 
triangular and filling the emarginations of segment 8. Genital organs 
themselves quite simple; paramere two short, freely articulate lobes 
fitted onto the apex of the large troughlike basal piece which more or 
less envelops a simple, small, somewhat curved, and tubelike median 
lobe; no evidence of a tegminite. 

Affinities of the family. — The heteromerous tarsi place this family 
in the Tenebrionoidea; the open anterior coxal cavities place itjn^the 
group including the Oedemeridae, Pythidae, and Serropalpidae, as 
distinguished from the group including the Alleculidae,\Lagriidae, 
and Tenebrionidae, all of which have closed coxal cavities. The ser- 
rate claws with fleshy pulvilli and the laterally fused eighth abdominal 
segment of the male separate it from the others of this group of Teneb- 
rionoidea. The very small median lobe and the larger paramere 
with the small lateral lobes show affinities with the genus Mycterus of 



BEETLE FAMILY CEPHALOIDAE — ARNETT 



157 




U 



(^ 



158 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

the family Pythidae. In addition, Mycterus and Cephaloon both 
have prominent procoxae and mesocoxae and the prothorax is with- 
out lateral margins. Most workers in the past have considered the 
Oedemeridae and Cephaloidae closely allied, but my incomplete 
studies of the male genitalia of the heteromerous beetles indicate that 
this is not so. The very different structure of the apical abdominal 
segments of the species of Cephaloon leads me to suppose that these 
species are far removed from the oedemerids, but their true affinities 
are not yet recognized. 

Notes on the distribution oj the species. — As can be seen from the 
accompanying map (fig. 20), the species of this family show the typical 
distribution pattern of a Holarctic group which was probably derived 
from some Asian stock and spread to North America via the Alaskan 
land bridge. It is interesting to note that, based on both external 
morphology and the morphology of the male genitalia (in the case 
of C. pollens) both of the Asian species are most similar to two of the 
Eastern North American species and not to the Western North 
American species. If these similarities of morphology reflect relation- 
ship, as wo assume they do, then we again see the often-repeated 
pattern of a pre-glacial distribution across northern Canada from 
Alaska, down through Ontario and into Eastern United States. 
This I believe to be a further stock-piling of evidence against the 
theories of continental drift and the North Atlantic land bridge idea. 

Genus Cephaloon Newman 

Ichnodes Dejean, 1834, Cat. Col. ed. 3, p. 227. {Nomen nudum, one ms. trivial 
name listed.) 

Cephaloon Newman, Ent. Mag., vol. 4, p. 376, 1838. (Genotype, Cephaloon 
lepturides Newman; 1838; monobasic.) 

Cephalaon Motschulsky in Schrenck, Reisen und Forschungen in Amurlande, 
vol. 2, pt. 2, p. 140, I860. (Error for Cephaloon.) 

Typitium Casey, Ent. News, vol. 9, p. 193, 1898. (Genotype, Cephaloon ungulate 
LeConte, Proc. Boston Soc. Nat. Hist., vol. 16, p. 275, 1873; original desig- 
nation and monobasic. Placed in synonymy by Hopping and Hopping, 
Pan-Pacific Ent., vol. 10, p. 64, 1934.) 

Sponidium Casey, Ent. News, vol. 8, p. 193, 1898. (Genotype, Cephaloon 
tenuicorne LeConte, 1873; present designation. Placed in synonymy by 
Hopping and Hopping, 1934.) 

Ephamillus Semenow, Horae Soc. Ent. Ross., vol. 34, p. 495, 1900. (New 
synonymy. Genotype, Cephaloon variabile Semenow «= Cephaloon variabilis 
Motschulsky in Schrenk, 1860; original designation and monobasic.) 

Drachylis Casey, 1898, Ent. News, vol. 9, p. 195. (New synonymy. Genotype, 
Drachylis simulans Casey, 1898; monobasic.) 

Discussion. — The type and only known specimen of Drachylis 
simulans Casey, the genotype of Drachylis Casey, has been examined. 
This is an unfortunate example of the description of a species on one 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 103. PLATE 5 




Adult, male genitalia, and claws of Cephaloidae: 1, Cephaloon ienuicorne LeCorite, female, 
showing general habitus; 2, paramere of male, C. tenuicorne; 3, paramere of male, C. 
bicolor Horn; 4, paramere of male, C. pacificum Van Dyke; 5, paramere of male, C. 
lepturides Newman; 6, apical portion of paramere of male, C. ungulate LeConte; 7 
abdominal segments 7-9 of male, C. ungulare; 8, median lobe of male, C. pallens 
Motschulsky; 9, median lobe of male, C. lepturides Newman; 10, hind claw of C. tenui' 
come; 11, hind claw of C. pallens; 12, hind claw of C. variabilis Motschulsky. 



BEETLE FAMILY CEPHALOIDAE — ARNETT 159 

poor specimen. Casey was certainly justified in wanting to describe 
this specimen for he beheved that it lacked the comblil\:e claws charac- 
teristic of the other species in the family, and therefore, its description, 
even though based on a single specimen, would alert collectors to hunt 
for such a strange cephaloid. A close examination, however, reveals it 
to have been patched, the legs being undoubtedly from some other 
beetle. It appears to be Cephaloon bicolor Horn, with which I synony- 
mize it. 

The genus Ephamillus Semenow is based ^; on the same variable 
characters possessed by the other genera erected for species of this 
family. K6no (Fauna Nipponica, vol. 10, fasc. 8, No. 10, pp. 76-82, 
1937) illustrates three characters, which, if they were constant and as 
illustrated, would serve for recognition of a genus. However, none of 
them appears to be constant or as distinctive as thought by Kono and 
others. The acute pulvilli, curved at the tips, are found in three 
species, C. jpacificum, C. ungulare, and C. variabilis. The shape of the 
pronotum, as illustrated by Kono for C. variabilis, is subject to the 
same sort of variation in all the species. Finally, the sinuation of the 
hind tibia of G. variabilis, reported to be so pronounced that a portion 
of the tibia is thrown out of line at least a distance equal to the width 
of the tibia, is often barely perceptible even under a microscope, and 
on some specimens of the species it cannot be seen at all. For these 
reasons, I feel that this genus is invalid and I place it in synonymy 
with Cephaloon. 

The marginate pronotum eliminates the genus Stenocephaloon Pic, 
1932 (Melanges Exotico-Entomologiques, fasc. 59, p. 2; genotype, 
Stenocephaloon metallicum Pic, monobasic) from this family. Until 
specimens can be studied, it is best placed in the family Serropalpidae 
near the genus Stenotrachelus Berthold. 

The following key to the known species of this family is adapted 
from Hopping and Hopping (Pan-Pacific Ent., vol. 10, pp. 64-70, 
1934) . For identification purposes, the illustrations which accompany 
that paper are very useful. 

Key to the species of Cephaloidae 

1. Pulvilli of tarsal claws slender, acute, and curved at tips (pi. 5, fig. 12) 2 

Pulvilli of tarsal claws robust, obtuse, not curved at tips (pi. 5, figs. 10, 11) _ 4 

2. Three distal antenoal segments together approximately 3 mm. long; hind 

femora at most simply curved; 1 to 15 mm. (United States) 3 

Three distal antennal segments together not over 1.5 mm. long; hind femora 
tending to be sinuate; size 17 to 20 mm. (Amur River, Eastern Siberia, and 
Japan) C. variabilis Motschulsky 



160 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

3. Lateral margins'of pronotum behind middle_ distinctly [emarginate; Western 

United States C. pacificum VanDyke 

Lateral margins of pronotum behind middle almost straight (Eastern United 
States) C. ungulare LeConte 

4. Antennae with last three segments thickened 5 

Antennae with last three segments not markedly thickened 6 

5. Elytra without sutural and marginal black stripes (Eastern United States) . 

C lepturides Newman 

Elytra with sutural and marginal black stripes (Amur River, Eastern Siberia 

and Japan) C. pallens Motschulsky 

6. Three distal antennal segments together approximately 1.5 mm. long (West 

Coast of United States) C. bicolor Horn 

Three distal antennal segments together barely over 1 mm. long (Western 
United States and Canada) 7 

7. Male with bifurcate process of third visible abdominal sternite long and finger- 

like; female with apical abdominal sternite shallowly and narrowlj' emargi- 

nate; color of female black to testaceous C. tenuicorne LeConte 

Male with bifurcate process of third visible abdominal sternite shorter, more 
triangular; female with apical abdominal sternite deeply emarginate; color 
of female reddish testaceous with elj'tra and metasternum black. 

C. vandykei Hopping and Hopping 

Key to the species of Cephaloon, based upon the male genitalia 

Male of Cephaloon variabilis Motschulsky unknown. 

1. Ratio of length of paramere to length of basal piece not over 1:2 2 

Ratio of length of paramere to length of basal piece not less than 1:3.5 3 

2. Median lobe tapering abruptly at apical third, slender (pi. 5, fig. 8) ; paramere 

lobes long and slender C. pallens Motschulsky 

Median lobe evenly tapering from base, more robust (pi. 5, fig. 9) ; paramere 
lobes shorter and heavier (pi. 5, fig. 5) C. lepturides Newman 

3. Paramere lobes acute at apex (pi. 5, fig. 6); median lobe long and slender, 

sides more parallel C. ungulare LeConte 

Paramere lobes enlarged at apex; median lobe shorter and more robust at 
apex 4 

4. Apex of paramere lobes about twice width of base when viewed laterally 

(pi. 5, fig. 3) C vandykei Hopping and Hopping and C. bicolor Horn 

Apex of paramere lobes about same width as at base 5 

5. Apex of basal piece laterally extending beyond base of paramere to about 

one-half length of paramere lobes; paramere lobes stout, blunt at apex, 

uniform in width (pi. 5, fig. 4) C, pacificum VanDyke 

Apex of basal piece laterally extending only slightly beyond base of paramere; 
paramere lobes more slender, and slightly enlarged at apex (pi. 5, fig. 2). 

C. tenuicorne LeConte 

In view of the relatively recent and thorough discussion of the 
species by Hopping and Hopping (1934), I feel that it is superfluous 
to repeat it here except to mention that under C. bicolor the name 
Drachylis simulans Casey should be added as a synonym (see discus- 
sion under generic synonymy preceding) . 



BEETLE FAMILY CEPHALOIDAE — AENETT 161 

There follows the bibliographic citations of the two exotic species 
not mentioned by Hopping and Hopping. The key and illustrations, 
I believe, sufficiently characterize them. A revised section of the 
Coleopterorum Catalogus (Junk) is being prepared which will give all 
of the literature references, therefore they will not be cited here. 

Cephaloon pallens Motschulsky, 1860 

Plate 5, Figures 8, 11 

Cephaloon pallens Motschulsky, in Schrenck, Reisen und Forschungen in Amur- 
lande, vol. 2, pt. 2, p. 140, pi. 9, fig. 15, 1860. (Type locality, Kisi, on the 
River Amur.) 

Cephaloon pallens var. cindipennie Heyden, Deutsche Ent. Zeitschr., p. 167. 
1892. (Type locality unknown, except "Amur" as given in the title of the 
paper.) 

Cephaloon pallens var. koltzei Heyden, Deutsche Ent. Zeitschr., p. 168, 1892. 
(Type locality, "Amur," as above.) 

Cephaloon pallens var. maculicolle Heyden, Deutsche Ent. Zeitschr., p. 167, 1892. 
(Type locality, "Amur," as above.) 

Cephaloon pallens var. picticolle Heyden, Deutsche Ent. Zeitschr., p. 167, 1892. 
(Type locality, "Amur," as above.) 

1 Cephaloon pallens Motschulsky, Solsky, Horae Soc. Ent. Rossicae, vol. 11, p. 
295, 1875. (Djalinda River. Note: This is probably a misidentification of 
Ephamillus variabilis var. tristiculus Heyden, 1892, judging from the descrip- 
tion and locality.) 

Cephaloon sakurae Lewis, Ann. Mag. Nat. Hist., ser. 6, vol. 15, p. 444, fig. 10, 
1895. (Placed as synonym by K6no, 1937, Fauna Nipponica, vol. 10, fasc. 
8, No. 10, p. 79.) 

In a species with the extreme color variation exhibited in all of the 
species of this family, the naming of a few of the color variants serves 
no useful purpose and these are therefore here disregarded. 

Cephaloon variabilis Motschulsky, 1860 

Plate 5, Figure 12 

Cephaloon variabilis Motschulsky, in Schrenck, Reisen und Forschungen in Amur- 
lande, vol. 2, pt. 2, p. 141, 1860. (Type locality, Marunsk on the River 
Amur.) 

Cephaloon variabilis var. tristiculus Heyden, Deutsche Ent. Zeitschr., 1892, p. 169. 
(Type locality, unknown except "Amur.") 

Here again, the naming of a single color form can serve no useful 
purpose. 



II, S. GOVERNMENT PRINTING OFFICE: 1953 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 




SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 



Vol. 103 Washington : 1953 No. 3322 



THE FRESH-WATER TRICLADS (TURBELLARIA) OF ALASKA 



By Roman Kenk 



Introduction 



In 1948, during an investigation of biting insects in Alaska con- 
ducted by the Bureau of Entomology and Plant Quarantine, United 
States Department of Agriculture, Dr. Reese I. SaUer collected and 
transmitted to me several samples of fresh-water triclads (planarians) . 
An examination of the material revealed that the worms belonged to 
the genus Polycelis, a genus fairly common in Asia and Europe but 
only twice reported from North America. The finding suggested a 
closer relationship of the Alaskan fresh-water fauna with that of East 
Asia. A more thorough study of the triclads of Alaska promised to 
yield a more definite understanding of these zoogeographical relations. 

My field trip ^ to Alaska was made in the sunmaer of 1950. Since 
the available time was rather limited, the collecting was done mainly 
along the highways of the territory and in the vicinities of Point 
Barrow and Umiat. 

The following roads and fresh-water localities were visited on the 
trip: Steese Highway in the section between Fairbanks and Birch 
Creek (milepost 103); Elliot Highway, from Fairbanks to Livengood; 
tundra lakes and pools in the vicinity of Point Barrow; ColvUle 
River, several streams, and a lake near Umiat; Mount McKinley 
Park Road, a section of about 15 miles adjoining the railroad station; 
Glenn Highway, from Anchorage to Glenallen; road from Palmer to 

' This study was supported by the Arctic Institute of North America under contractual arrangement 
with the OfBce of Naval Research. 

238538—53 1 163 



164 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Willow; road from Anchorage to Potter; and Richardson Highway, 
section between Valdez and Glenallen. 

No fresh-water triclads were found by me either at Point Barrow 
or at Umiat. I received, however, from the U. S. National Museum, 
a sample of triclads collected by P. F. Scholander in a lake near Umiat. 
The relative scarcity of planarians in tundra lakes and pools may be 
due to the high acidity of the waters or to the extremely severe con- 
ditions prevailing in them during the long winter season. 

Grateful acknowledgment is made of the very helpful cooperation 
which was extended to me by several agencies in Alaska: The Arctic 
Research Laboratory, Point Barrow; the U. S. Public Health Service, 
Anchorage; the Alaska Road Commission, Fairbanks, Anchorage, and 
Glenallen; and the Division of Forestry, Bureau of Land Management, 
Fairbanks and Anchorage. I also wish to express my indebtedness to 
Prof. Edward G. Reinhard of the Catholic University of America and 
to Dr. Fenner A. Chace, Jr., and Dr. Doris M. Cochran of the U. S. 
National Museum who kindly permitted me to use their laboratory 
and office facilities in Washington, D. C. 

Four species of fresh-water triclads were collected in Alaska. Two 
are inhabitants of the White Mountains, a mountain range extending 
in an east-west direction north of Fairbanks. The other two are 
widely distributed in waters of the Alaska Range and of the southern 
section of Alaska and one of them reaches as far north as Umiat. 

Family Planariidae 
Genus Phagocata Leidy 

Phagocata niveau new species 

Figure 21; Plate 6, Figure 1 

Description. — This is a slender, rather delicate species. Mature 
specimens measure up to 8 mm. in length and about L5 mm. in width. 
In the quietly gliding animal the anterior end is truncated,with a very 
slightly bulging frontal outline and with rounded lateral corners 
(auricles) . There is no distinct narrowing or neck behind the auricles 
and the lateral margins of the head are approximately parallel. Behind 
the head, the body widens and soon reaches its greatest width. From 
there on, the lateral margins of the body run parallel up to the level 
of the mouth, to converge agam in the postpharyngeal region and to 
meet in a bluntly pointed posterior end. 

The species lacks pigment and usually appears white and somewhat 
transparent. The intestinal contents may shine through the body 
wall and give the animal a certain amount of color; the margins of the 
body, the head region, and the areas occupied by the pharynx and the 
copulatory apparatus, however, are always white. 



FRESH-WATER TRICLADS OF ALASKA — ^KENK 165 

There are two rather small eyes, situated close together (about one- 
fourth the body width apart at the level of the eyes) and far removed 
from the frontal end. This character, easily recognized in life, dis- 
tinguishes the species from another white triclad with which it shares 
its habitat, Dendrocoelopsis alaskensis, described as new on p. 178. 

The pharynx is inserted, in sexually mature specimens, somewhat 
behind the middle of the body and measures about one-sixth of the 
body length. The copulatory organs occupy the anterior half of the 
postpharyngeal region. 

The animal moves by gliding only; crawling movements, such as 
are seen in other triclads, particularly in those equipped with anterior 
adhesive organs, have not been observed in this species. 

From the description it may be seen that the species in life shows 
a close resemblance to other species of the same genus, particularly 
to the American Phagocata morgani, the European P. albissima, P. 
vitta, and related forms. A separation of these species can be made 
only on the basis of anatomical characters. 

Only those characters of the digestive system that have a taxonomic 
significance are discussed here. The pharynx has a structure typical 
of the family Planariidae; i. e., the fibers of the internal muscle zone 
are arranged in two distinct layers, a thick inner circular layer and a 
narrower outer longitudinal one. The anterior intestinal trunk bears 
10 or 11 branches on each side. Each posterior trunk has 21 to 27 
lateral branches and numerous short medial branches in both the 
pharyngeal and postpharyngeal regions. 

The testes are numerous and are arranged, on each side of the 
midline, in a longitudinal zone extending from a short distance behind 
the ovary almost to the posterior end of the body. Their position is 
predominantly ventral, below the intestinal branches. Only a few 
testes extend into the mesenchymatic "septa" between the branches 
toward the dorsal side. 

The two ovaries are typical, each situated approximately below the 
second intestinal branch. An undifferentiated mass of cells, the par- 
ovarium, is attached to the dorsolateral side of each ovary. 

The genital pore (pg), situated about halfway between the mouth 
and the posterior end of the body, leads into a small cavity (ac) 
which continues to the left and dorsally into the duct of the copulatory 
bursa (bd) and to the right and anteriorly into the male atrium (am) . 
This cavity may be considered to represent a common genital atrium. 
In some specimens, however, there appears to be no differentiation 
of the atrium into male and common parts, and the bursa duct and 
an undivided atrium meet at or near the genital pore. These varia- 
tions are obviously due to the different states of muscular contraction 
in which the animals were killed. The atrium, narrow at the genital 
aperture, widens as it extends forward, to the right side of the midline. 



166 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

It is lined with a tall, glandular epithelium, the cells of which project 
into the cavity in a villuslike fashion. Below the epithelium there 
are two layers of muscle fibers, one circular and the other longitudinal. 

The penis consists of a spherical, muscular bulb embedded in the 
mesenchyme, and a moderately large papilla projecting into the male 
atrium. The bulb is pierced by numerous gland ducts which open 
into the lumen of both the bulb and the papilla. The shape of the 
papilla is subject to great variation, due apparently to the state of 
contraction of the organ. It may be twisted to one side and even 
partly inverted into the lumen of the penis (similar to the pseudo- 
flagellum of various dendrocoelids) . The shape shown in figure 21 
appears to be that of the organ at rest. The outer wall of the papilla 
is covered with a tall to cubical epithelium similar to that lining the 
atrium. Below the epithelium there is a layer of circular muscle 
fibers followed by another of longitudinal fibers. The shape of the 
penis lumen {Ip) is as changeable as that of the papilla. Typically, 
it appears to be wider in the bulb than it is in the papilla, though 
there is no distinct ejaculatory duct differentiated. The lumen opens 
ventrally to the tip of the papUla. The two vasa deferentia {vd) 
penetrate the penis bulb from both sides and empty into the penis 
lumen separately, but not far apart. 

The two oviducts converge at the level of the copulatory apparatus, 
the left one passing between the bursa duct and the male atrium, and 
unite at a point dorsally to the atrium. The rather long common 
oviduct {ode) curves ventrally and opens into the posterior part of the 
male atrium. The terminal sections of the paired oviducts and the 
greater part of the common oviduct receive the outlets of numerous 
eosinophilic glands, the cell bodies of which are scattered in the sur- 
rounding mesenchyme, particularly dorsally to the atrium. 

The copulatory bursa (6) is of moderate to large size and is ir- 
regularly lobed. The bursa duct or stalk {hd) is wide, runs posteriorly 
to the left of the midline, and curves ventrally to reach the genital 
aperture. It is lined with a tall, glandular epithelium and surrounded 
with a strong muscular coat consisting of intermingled circular and 
longitudinal fibers. 

Taxonomic position. — The genus Phagocata Leidy (Fonticola 
Komarek) in its present extent (cf. Hyman, 1937, pp. 300-302) has 
representatives in Europe, Asia, and North America. The genus is 
not quite homogeneous and will probably, in due time, be subdivided 
into several genera (cf. Beauchamp, 1939). For the purpose of com- 
parison, we may consider here only those species of the genus that 
lack pigment. Though the presence, or the lack, of pigment is a 
character of subordinate taxonomic value, it may nevertheless serve 
well as a character of specific rank. The Alaskan form differs from 



FRESH-WATER TRICLADS OF ALASKA — ^KENK 



167 



European species such as P. vitta (Duges) and P. albissima (Vejdovsk^) 
and from the American white P. morgani (Stevens and Boring) mainly 
in the structm-e of the male copulatory organ. Asia has several 
species of Phagocata, the majority of them pigmented forms. Of the 
three unpigmented Asiatic species that may belong to the genus, two 




Figure 21. — Phagocata nivea, diagram ot the copulatory organs In longitudinal section, 
X 92. ac, common atrium; am, male atrium; b, copulatory bursa; bd, bursa 
stalk; Ip, penis lumen; o, mouth; ode, common oviduct; pg, genital pore; vd, vas deferens. 



have been described from immature specimens and the anatomy of 
their reproductive systems is not known: Planaria pellucida Ijima 
and Kaburaki (1916) from Sakhalin and a species from the Bailial 
region assigned tentatively to Fonticola by Bazikalova (1947). A 
third species, Phagocata coarctata (Arndt, 1922), from the vicinity of 
Vladivostok, is sufficiently well Imown, although no fully mature 
individuals of this species have been studied. P. coarctata differs 
from P. nivea externally in being smaller and broader, and in having 
a different contour of the anterior end, which bears protruding lateral 
lobes, and a greater distance between the two eyes. Anatomically, 
the two species are, undoubtedly, closely related.^ 



' Livanov and Zabusova (1940, p. 146) state that a reexamination of Amdt's slides of PUmaria coarctata 
showed that the arrangement of the muscle fibers of the pharynx conformed with the dendrocoelid type 
(circular and longitudinal fibers of the internal muscle zone intermingled). Amdt (1922, p. 108) described 
the anatomy of the pharynx in minute detail and indicated, both in a figure (pi. 4, fig. 7) and in the text, 
a typical planariid pattern. I must assume that some confusion occurred somewhere, probably in the 
identification of the slides sent to Livanov. 



168 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

In Phagocata nivea the penis lumen opens usually below the tip of 
the penis papilla. This character has been used by Livanov and 
Zabusova (1940, p. 96) to segregate a group of Asiatic species from 
Phagocata and to place them in a new genus, Penecurva. The same 
character is found, however, in a common North American species, 
Phagocata morgani, which shows no other close relations with the 
Asiatic group. This character is apparently inadequate as a basis 
for the establishment of a new genus. 

Holotype. — On one slide, USNM 22332, creek crossing EUiot High- 
way at milepost ^31.0, July 24, 1950. 

Distribution and ecology. — Phagocata nivea was collected in cool, 
fast mountain streams on the slopes of the White Mountains, a range 
north of Fairbanks. The temperature of the water ranged from 3.2 
to 7.2° C. (July). The animals are cold-stenothermic and do not 
tolerate sudden increases in temperature. They were often found 
in the company of another white triclad, Dendrocoelopsis alaskensis. 

Stream on Steese Highway (pi. 8), at milepost 84.0, altitude 2,700 feet, July 
19 and 21, 1950, water temperature 6.9° C; one immature and one mature speci- 
men, from under stones. 

Willow Creek, on Steese Highway, at milepost 96.6, altitude 2,100 feet, July 
19, 1950, 7.2° C; two immature specimens, from under stones. 

Spring and stream on Steese Highway, milepost 82.5, near Alaska Road Com- 
mission camp, July 21, 1950; two mature and six immature specimens, on the 
undersides of stones. 

Fox Gulch, on Elliot Highway, milepost 1.2, July 24, 1950, 6.0° C, one small 
specimen near bait (beef liver). 

Creek crossing Elliot Highway at milepost 31.0, July 24, 1950; 39 specimens, 
about half of them mature, collected under stones (holotype). 

Genus Polycelis Ehrenberg 
Polycelis borealis^ new species 

Figure 22; Plate 6, Figure 2 

Description. — Mature animals are usually 12 to 15 mm. long and 
1.5 to 2 mm. wide (larger specimens, measuring up to 20 mm. in 
length, have been seen). The frontal margin is slightly convex. At 
the lateral corners of the head there is a pair of elongated, bluntly 
pointed auricles which are held raised when the animal is gliding 
quietly. Behind the auricles, the body first narrows slightly, then 
gradually widens, reaching its greatest width at the level of the 
pharynx. Behind the pharynx the body tapers to a moderately 
pointed posterior end. 

The color of the dorsal side is usually a uniform light or dark 
brown, that of the ventral side a light grayish brown. In animals 

' The large highways of Alaska are marked with wooden mileposts indlcatmg only full miles. Many 
posts were missing at the time when the collections were made. Fractional milages were usually estimated 
from the nearest milepost or from the speedometer readings of the car used. 



FRESH-WATER TRICLADS OF ALASKA — ^KENK 169 

from some localities, an indistinct lighter median line occurred dor- 
sally in the prepharyngeal region, with lighter areas above the pharynx 
and the copulatory organs. 

The species has many small eyes (a generic character) arranged 
in a band along the frontal margin of the head, the base of the auricles, 
and the lateral margins of the body a short distance behind the head. 
Anteriorly the eyes are placed in more than one row, somewhat 
irregularly scattered ; behind the head they are in a single row reaching 
backward about one-third to one-half the length of the prepharyngeal 
region. There is no narrowing of the band of eyes at the base of the 
auricles. 

The pharynx is inserted at about the middle of the body and 
measures in length almost one-fourth the length of the body. The 
copulatory organs occupy more than half the post-pharyngeal region. 

The animal moves by gliding only. 

The pharynx of Polycelis borealis is structurally typical of the genus 
Polycelis and of the family Planariidae, the muscle zone being formed 
by two distinct layers, a thick circular layer adjoining the epithelium 
of the pharyngeal lumen and a thinner layer of longitudinal fibers. 
The anterior trunk of the intestine bears 5 to 6 lateral branches. 

The numerous, fairly large testes are arranged in two zones, to 
the right and left of the anterior intestinal trunk, extending from the 
level of the ovaries posteriorly to the base of the pharynx. The 
testes are essentially ventral, though individual vesticles may extend 
dorsally in the mesenchymatic spaces between the intestinal branches 
and occupy almost the entire dorsoventral diameter of the body. 
Where intestinal branches are present, however, the testes develop 
only below them. Each zone of testes reaches laterally only little 
beyond the ventral nerve cord. 

The ovaries are situated behind the first pair of lateral branches of 
the anterior trunk of the intestine. 

The genital pore (j)g), situated in the midline behind the middle of 
the postpharyngeal region, connects with a narrow posterior extension 
of the genital atrium (a) . This extension leads dorsally and somewhat 
to the left into the duct of the copulatory bursa and anteriorly into 
the widened portion of the atrium containing the penis. There is no 
marked division of the atrium into a posterior common atrium and 
an anterior male atrium. The wall of the atrium is lined with a 
cubical epithelium and equipped with two muscle layers, one circular 
and the other longitudinal. 

The penis consists of a bulb embedded in the mesenchyme and a 
papUla projecting into the atrium. Neither of the two parts is very 
muscular. The shape of the penis, particularly of the papilla, is very 



170 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i03 

changeable. In specimens that were fixed in a well-extended condi- 
tion, the bulb appears spherical and the papilla has a conical shape. 
The bulb contains a spacious, irregularly lobed cavity, the seminal 
vesicle (vs). Numerous gland ducts empty into the lumen of the 
bulb after entering the bulb, particularly from its anterior surface, 
and penetrating between its muscle fibers. The ducts are filled 
with a granular, slightly eosinophilic secretion. The two vasa 
deferentia (vd) open into the seminal vesicle separately from the 
anterolateral sides. 

The penis papUla is conical or finger-shaped when well extended. It 
is covered with a cubical epithelium below which there are two rather 
feebly developed muscle layers, tapering in thickness toward the tip 
of the papilla: a layer of circular fibers adjoining the epithelium and 
a thinner layer of longitudinal ifibers. The absence of a strong mus- 
cular wall probably accounts for the great variation in the shape of 
the penis papilla observed in the material. The lumen of the seminal 
vesicle continues into the papilla as a wide canal (de), opening at or 
near (ventrally to) the tip of the papUla. The canal is lined with an 
epithelium of cubical cells and lacks the gland openings character- 
istic of the seminal vesicle. It corresponds to an ejaculatory duct 
but it apparently has no distinct muscle coat. 

The two oviducts, riuming posteriorly along the ventral nerve 
cords, bend dorsally and medially at the level of the copulatory 
organs. The left one passes through the space between the atrium 
and the bursa duct and unites with the oviduct of the right side 
dorsally to the atrium. The common oviduct {ode) runs ventrally 
and opens into the posterior part of the atrial cavity. The terminal 
sections of the paired oviducts and a section of the common oviduct 
are equipped with strongly eosinophilic shell glands. 

The copulatory bursa (6) is a large sac with a lobed outline. It 
connects posteriorly with a duct of almost uniformly wide diameter, 
the bursa stalk (bd). The duct runs first posteriorly, to the left of 
the penis bulb, then curves ventrally and opens, from the dorsal side, 
into the narrow terminal portion of the atrium, close to the genital 
pore. The bursa stalk has a strong coat of intermingled circular and 
longitudinal muscle fibers. There is no histological differentiation 
into anterior and posterior sections of the bursa stalk. 

Taxonomic position. — Polycelis borealis is the second species of the 
genus Polycelis to be found on the North American continent. The 
other species, P. coronata (Girard), reported from Wyoming and South 
Dakota (Hyman, 1931), resembles it closely in external appearance 
and probably cannot be distinguished from it in life. In both species, 
the shape of the anterior end is very similar. The auricles of P. 
borealis are perhaps a trifle longer and more pointed than those of P. 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 103, PLATE 6 






Sketches of living specimens of the four Alaskan triclad species, all XIO: 1. Phagocata 
nivea; 2. Polycelis borealis; 3. Dendrocoelopsis piriformis, striped form; 4. Dendrocoelop- 
sis alaskensis. 



FRESH-WATER TRICLADS OF ALASKA — KENK 



171 



coronata. Hyman (1931, p. 126) states that in P. coronata the 
curved band of eyes narrows as it crosses the base of the auricles; 
no such narrowing has been seen in P. borealis. These differences 
are, however, insignificant and do not permit a clear separation of 
the two forms. The same may be said of other species of the genus, 
which likewise could be confused with the American forms in life: 
the Japanese species, P. auriculata and P. karafto; and P. schmidti 
occurring both in Kamchatka and in Japan. These forms differ 
from each other more clearly in their anatomical characters. 

bd ode 




Figure 22. — Polycelis borealis, diagram of the copulatory organs in longitudinal section, 
X 70. a, atrium; b, bursa; bd, bursa stalk; de, ejaculatory duct; o, mouth; ode, common 
oviduct; pg, genital pore; vd, vas deferens; vs, seminal vesicle. 

To gain a better insight into the systematic relations of Poly- 
celis borealis to other species of the same genus, it is necessary to review 
the present state of the systematics of the genus. The genus Poly- 
celis has a muscular pattern of the pharynx typical of the family 
Planariidae; the oviducts unite, without embracing the bursa stalk, 
to form an unpaired terminal oviduct; the testes are situated in the 
anterior part of the body only; the male atrium is not surrounded 
by radial muscle plates; and the eyes are numerous. The genus 
thus defined (Kenk, 1930) has today about twenty species. It may 
be subdivided into subgenera on the basis of structural characters 
of the copulatory apparatus. 

Subgenus Polycelis, lacking adenodactyls and lacking an excessive 
development of the muscle coat of the male atrium. This subgenus 
includes Sorocelides Sabussowa (1929, p. 521) and Polycelidia Zabu- 
sova (1936, p. 152), both described as distinct genera. The subgenus 
comprises the following species: 



238538—53 



172 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Polycelis nigra (O.F.MulleT), Europe P. polyopis Zabusova (1936), Kam- 

P. receptaculosa (Livanov and Zabusova, chatka 

1940), Teletskoe Lake in the Altai P. karafto Ijima and Kaburaki (1916), 
Mountains Sakhalin 

P. eburnea (Muth, 1912), Aral Sea P. sopporo (Ijima and Kaburaki, 1916), 
region Japan 

P. ttfeeftca Hyman (1934), Tibet P. coronata (Girard, 1894), Wyoming 

P. koslowi (Zabusov, 1911), Tibet and South Dakota 

P. elongata (Sabussowa, 1929), Kam- 
chatka 

Subgenus Seidlia Zabusov (1911, suppl., p. 7), distinguished by an 
extraordinarily thick muscle zone surrounding the male atrium. 
Zabusov's (1916, p. 273) genus Rjabuschinskya likewise belongs here. 
Species of the subgenus: 

Polycelis sabussowi (Seidl, 1911), in- P. schmidti (Zabusov, 1916), including 
eluding Seidl's species Sorocelis P. ijimai Kaburaki (1922), Kam- 

sabussowi, S. lactea, S. stummeri chatka, Kurile Islands, and Japan 

(cf. Kenk, 1936), Turkestan P. auncu/a<a Ijima and Kaburaki (1916), 

P. relicta (Sabussowa, 1929), Kamchatka Japan 

P. eurantron (Zabusova, 1936), Kam- 
chatka 

The third subgenus, Ijimia Bergendal (1890, p. 326), is charac- 
terized by possessing solid adenodactyls : 

Polycelis felina (Dalyell), including P. linkoi Zabusov (1901), Onega Lake 
P. cornuta Johnson, Europe and in European Russia 

North Africa P. oculi-marginata (Palombi, 1931), New 

P. tenuis Ijima, Europe and western Guinea (see Beauchamp, 1947) 

Asia 

Two species of the genus Polycelis are too mcompletely known to 
permit their assignment to either the subgenus Polycelis or Seidlia: 
P. eudendrocoeloides (Sabussowa, 1929) from the Kamchatka Penin- 
sula and P. tibetica (Zabusov, 1911, p. 349) from Tibet (P. tibetica 
Hyman, 1934, will have to be renamed if it should prove to be different 
from Zabusov's species) . 

Polycelis borealis is clearly a member of the subgenus Polycelis. 
It differs from P. nigra in having weU-developed auricles; from P. 
elongata, P. polyopis, and P. sapporo, in the arrangement of the eyes; 
and from P. coronata and P. receptaculosa in the structure of the male 
copulatory organ. The reproductive system of P. karafto has not been 
described adequately and thus does not permit a comparison with 
that of the Alaskan species. The remaining species of the subgenus, 
P. eburnea, P. koslowi, and P. tibetica Hyman, two of which are known 
only from the study of preserved specimens, agree with P. borealis 
in being pigmented forms with prominent auricles and in having 
a similar anatomy of the copulatory organs. The four species are 



FRESH-WATER TRICLADS OF ALASKA — KENK 173 

undoubtedly very closely related; whether some of them are identical 
or are the same as the Alaskan form cannot be established on the basis 
of present knowledge. The Alaskan Poly cells, therefore, is described 
as a new species, 

Holotype. — On three slides, USNM 22333, clear spring on the road 
from Palmer to Willow, 20.1 mUes from Palmer, altitude 3,800 feet, 
Aug. 9, 1950. 

Distribution and ecology. — Polycelis borealis is a very common species 
occurring in mountain streams in the southern part of Alaska (Alaska 
Range, Talkeetna Mountains, Chugach Range). Typically it 
inhabits clear, cold, fast-running waters. It has also been found, 
however, in several small mountain lakes which connect with streams 
in which the species lives. It has not been observed in sUt-bearing 
glacier streams. 

The typical temperature range of the habitats was between 3.0° 
and about 15° C. (August 1950). Temperatures above 15° C. (up 
to 22.4° C.) were encountered only rarely and only in habitats which 
presumably have great diurnal temperature amplitudes (shallow lakes, 
small exposed streams). 

The great majority of the specimens collected was asexual. Sexually 
mature animals were seen in only 6 of the 31 localities in which the 
species was found. In some localities, a large percentage of the speci- 
mens exhibited freshly regenerated heads or posterior ends, indicating 
that vivid asexual reproduction was taking place. 

Clear springs on the road from Palmer to Willow, 20.1 miles from Palmer, 
altitude 3,800 feet, Aug. 9, 1950, water temperature 3.0° C, numerous specimens, 
five mature (holotype). 

Clear, fast mountain stream crossing Mount McKinley Park road, 5.5 miles 
from the railroad station; Aug. 5, 1950. 

Streams on road from Palmer to Willow 11.9, 18.0, 19.2, 20.2, 21.6, 22.1, 22.8, 
and 34.0 miles from Palmer, and Ice Lake, 20.4 miles from Palmer, Aug. 9, 1950. 

Streams on Glenn Highway (Anchorage to junction with Richardson Highway 
near Glenallen) at mileposts 33.1, 33.9, 43.5, 64.8, 89.4, 98.8, and 117.1; Aug. 11 
and 13, 1950. 

Streams on road from Anchorage to Potter 2.9 (Campbell Creek), 10.3, and 10.6 
miles from the city limits of Anchorage, Aug. 12, 1950. 

Streams on Richardson Highway (section between Valdez and Glenallen) at 
mileposts 20.5, 25.5 (22.4° C), 26.3 (see pi. 7), 29.0, 37.3, 54.7, 62.6 (see pi. 7), 
81.0 (Squirrel Creek), and 87.5 (Rock Creek), and lake at milepost 27.3 (22.1° C), 
Aug. 15, 1950. 

Polycelis borealis was also identified in samples of triclads collected 
by Dr. Reece I. Sailer in July and September 1948. The following 
localities were represented: Richardson Highway, mileposts 8.3, 187.5, 
192.9, 209.7, 223.6 and 239.9, and Glenn Highway, milepost 117.1. 



174 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Family Dendrocoelidae 
Genus Dendrocoelopsis Kenk 

Dendrocoelopsis piriformis, new species 

Figure 23; Plate 6, Figure 3 

Description. — Sexually mature specimens of this rather broad and 
plump species measure up to 15 mm. in length and 3 mm. in width. 
In quietly gliding animals, the head is truncated, with a convex 
frontal margin (grasping organ). There is a visible subterminal 
depression corresponding to the adhesive surface of the grasping 
organ or sucker. The rounded lateral corners of the head are formed 
by the auricles, which protrude only little laterally, thus causing an 
insignificant narrowing (neck) to appear behind them. Behind the 
head, the body margins widen gradually. The greatest width is 
reached at the level of the mouth. Behind that level, the body tapers 
again to a bluntly pointed posterior end. When the animal is at 
rest, the body appears short and wide, about pear-shaped, the body 
margins often forming a wavy or irregular contour. Yoimg specimens 
are more slender and their lateral margins are more nearly parallel 
when they are in gliding motion. At rest, however, they assume the 
same pyriform shape as the adults. In short, the habit of the species 
may be described as resembling that of European and Asiatic repre- 
sentatives of the genus Bdellocephala Man, with which it has in com- 
mon the broad shape and the anterior grasping organ. 

The two eyes are situated on the dorsal side of the head. Their 
distance from each other amounts to somewhat more than one-third 
the width of the head at the level of the eyes. The distance of each 
eye from the lateral margin is smaller than the distance from the 
frontal margin. 

The color of the dorsal side is usually a cloudy brown or dark 
brownish gray. In some specimens, the pigment is arranged in 
definite longitudinal stripes: one sharply marked median stripe flanked 
on each side by a light (usually yellow) band ; laterad to this band the 
body darkens, losing its pigment again near the lateral margin. In 
other specimens, lacking the two light dorsal bands, the back may be 
either uniformly pigmented or may show a more or less distinct darker 
band along the midline. Young specimens and striped adults clearly 
show the finer disposition of the pigment in small rounded dots, each 
dot apparently corresponding to an individual pigment cell. 

The pigment pattern of the dorsal side of the head is quite charac- 
teristic. A very dark field between the eyes extends anteriorly to 
both sides of the grasping organ. The area above the organ is unpig- 
mented and white, and so are the two elongated ocular areas antero- 



FRESH-WATER TRICLADS OF ALASKA — KENT 175 

laterad to the eyes. A pair of indistinct, converging streaks of lighter 
coloration may run posteriorly from the lateral angles of the head. 

The ventral side is light gray. 

The pharynx is situated, in mature animals, behind the middle of 
the body; the mouth, at the beginning of the last third of the body; 
and the genital aperture, midway between the mouth and the hind end. 

The normal locomotion of the animal is quiet gliding. When dis- 
turbed, it may attach itself to the substratum (apparently with the 
marginal adhesive zones) or move by "crawling." Young specimens 
do not crawl as readily as do adult ones. 

The grasping organ, or sucker, in living animals, appears as a well- 
marked bulge on the frontal margin of the head, showing a concave 
ventral depression. In preserved specimens, the frontal margin is 
generally curved ventrally and the site of the organ forms a thick, 
grooved rim. Anatomically, the organ consists of glandular and 
muscular elements. The subterminal adhesive surface is covered with 
an epithelium devoid of rhabdites and pierced by numerous gland 
ducts filled with a granular, eosinophilic secretion. The cell bodies 
of the glands are scattered through the mesenchyme of the anterior 
half of the prepharyngeal region, particularly above the intestine. 
The gland ducts run anteriorly in dense bundles and, in their terminal 
sections, are thickly swollen with secretion. The muscular system of 
the organ, which could not be analyzed in detail on account of the 
density of the glandular structures, has fibers attached to the adhesive 
surface, serving presumably as retractors. 

The rather short and thick pharynx is structurally typical of the 
family Dendrocoelidae; its internal muscular zone consists of inter- 
mingled circular and longitudinal fibers. 

Auricular sense organs are represented by two bands of sensory 
epithelium extending posteriorly from the sides of the frontal margin. 
The cells of this epithelium are less tall than are those of the sur- 
rounding body epithelum ; they lack rhabdites and are approached by 
nerve fibers from the underlying mesenchyme. The location of these 
organs corresponds to the two light, longitudinal streaks on the sides 
of the head seen in the living animal. 

The testes are of moderate size, numerous, densely packed, and 
occupy the dorsal half of the mesenchyme, generally above the intes- 
tinal branches. They are arranged in two wide areas on both sides 
of the midline, extending from the level of the ovaries close to the 
posterior end. 

The ovaries, situated at the level of the second pair of branches of 
the anterior intestinal trunk, show no structural peculiarities. 

The genital pore (pg) leads immediately into two cavities, the male 
atrium (am) and, to the right and somewhat posteriorly, the duct of 



176 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

the copiilatory bursa (bd). No common atrium is present, 
and both the papUla of the penis and the opening of the 
oviduct are situated in the "male" atrium. The atrium is a conical 
cavity, wide anteriorly and tapering toward the genital pore. It is 
lined with a cubical epithelium, below which occurs a thin layer of 
fine circular muscle fibers and a thicker layer of coarser longitudinal 
fibers. 

The penis consists of a spherical, muscular bulb and an elongated 
papilla. The penis bulb is differentiated into a wide peripheral 
muscular zone and a more central parenchymatic zone which contains 
a cavity, the seminal vesicle (vs) . The vesicle is lined with a glandu- 
lar epithelium and its wall forms villuslike projections. The two 
vasa deferentia (vd) penetrate the penis bulb from the anterolateral 
sides and open into the seminal vesicle on two prominent conical 
papillae projecting from the anterior wall of the vesicle a short dis- 
tance from each other. 

The papilla of the penis is finger-shaped, tapering toward the tip, 
and is highly muscular. It is covered with a thin cubical epithelium. 
Below the epithelium there is, in the basal portion of the papilla, a 
thin layer of longitudinal muscles, below which lie a stronger circular 
layer and a second longitudinal layer. In the distal part of the 
papUla, the external longitudinal layer is lacking. The axis of the 
papilla is pierced by the ejaculatory duct (de) which leads from the 
seminal vesicle to the tip of the papilla. The duct has a cubical 
epithelium and a strong coat of longitudinal muscle fibers. 

The two oviducts approach the midline in the region of the copula- 
tory apparatus, the right one passing between the atrium and the 
bursa stalk, and unite behind and above the atrium. The common 
oviduct (ode) proceeds ventrally, then curves anteriorly, and empties 
into the terminal part of the atrium close to the genital pore. 

The copulatory bm-sa (b) is a lobed sac lined with a tall glandular 
epithelium. The distal parts of the epithelial cells are filled with 
fine eosinophilic granules. The duct or stalk of the bursa (bd) is dif- 
ferentiated into a narrow anterior section with a weak muscular coat, 
which connects with the bursa, and a posterior wider section which 
bends ventrally and opens at the genital pore. The ceUs of the 
anterior section resemble those of the bursa in having similar granular 
inclusions. Those of the posterior section lack the inclusions and are 
ciliated. The muscle coat of the duct consists of intermingled longi- 
tudinal and circular fibers. 

Taxonomic position. — I have placed the species in the genus 
Dendrocoelopsis established originally for a European species, D. 
spinosipenis (Kenk). The original definition of the genus was based 
on the following characters: fibers of the inner muscle zone of the 



FRESH-WATER TRICLADS OF ALASKA — ^KENK 



177 



pharynx intermingled ; no adenodactyl ; penis papilla developed, penis 
bulb of simple structure; oviducts unite without embracing bursa 
stalk; zone of testes extending behind the level of the copulatory 
organs; anterior end with subterminal true sucker; eyes not numerous 
(Kenk, 1930). Subsequently Beauchamp (1932, p. 254) founded a 
new genus, Amyadenium, with two species, A. vandeli Beauchamp and 
A. brementi (Beauchamp), both from the Pyrenees. Two more 
species were reported by the same author in later papers, A. chattoni 
Beauchamp (1949, p. 60), again from the Pyrenees, and A. garmieri 
Beauchamp (1950, p. 65), from central France. Beauchamp recog- 
nized the close relation of the new genus to Dendrocoelopsis, but 




Figure 23. — Dendrocoelopsis piriformis, diagram of the copulatory organs in longitudinal 
section, X 60. am, male atrium; b, bursa; bd, bursa stalk; de, ejaculatory duct; o, mouth; 
ode, common oviduct; pg, genital pore; vd, vas deferens; vs, seminal vesicle. 

separated it from the latter on accoimt of the absence of a highly 
complex grasping organ, or true sucker (i. e., an adhesive organ sepa- 
rated from the surrounding mesenchyme by a muscle layer) . Hyman 
(1935) described a dendrocoelid species from Montana under the 
name of D. vaginatus. Again the main characters of the species 
coincide with those of D. spinosipenis with the exception of the 
grasping organ, which is of a simpler type. The new Alaskan species, 
D. piriformis, also falls clearly in the vicinity of the species enumer- 
ated, as does another species, D. alaskensis, the description of which 
follows on p. 178. Within this group of species there is a gradual 
differentiation of the grasping organ, which is absent in D. alaskensis, 
present as a moderately developed adhesive organ in D. vaginata, 
D. piriformis, A. brementi, A. vandeli, A. chattoni, and A. garmieri, 
and as a more highly developed and muscular sucker in D. spinosipenis. 
A similar wide variation of the structure of the anterior grasping organ 



178 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ids 

is seen in the genus Dendrocoelum Orsted where, however, true suckers 
are not known. In general, the taxonomic vahie of adhesive organs 
in triclads appears to be subordinated to that of other anatomical 
structures. I therefore tentatively modify the definition of the 
genus Dendrocoelopsis by omitting the presence of a sucker as a 
generic character, to include the species described as Dendrocoelopsis 
and Amyadenium. 

Dendrocoelopsis pirijormis differs from the other members of the 
genus (in its wider extent) in several characters: It is pigmented, 
whereas the others lack pigment and appear white; the presence of 
two eyes differentiates it from the three blind species, D. vandeli, D. 
brementi, and D. garmieri and from the many-eyed D. chattoni; and 
the dorsal position of the testes separates it from D. spinosipenis, D. 
vaginata, D. garmieri, and D. alaskensis. Apart from these most con- 
spicuous characters, the structure of the male copulatory organ of D. 
piriformis is distinctive. 

Holotype.— On five slides, USNM 22334, Moose Creek, on Glenn 
Highway, milepost 186, near Alaska Road Commission camp, Aug. 
U. 1950. 

Distribution and Ecology. — Dendrocoelopsis piriformis is a eury- 
thermic species and inhabits lakes and their outlets in the southern 
part of Alaska and also occurs in a lake near Umiat. 

Long Lake, on Glenn Highway, milepost 85.9, Aug. 11, 1950, clear water, 
17.8° C. (near bank) ; under stones, several specimens, two of them mature. 

Lake on Glenn Highway, milepost 88.1, Aug. 11, 1950, water temperature 
varying with depth (near bank, 19° C.) ; under stones, several specimens, one 
mature. 

Lake on Glenn Highway, milepost 23.5 (see pi. 8), Aug. 13, 1950, clear water, 
23.6° C. (near bank) ; several immature specimens. 

Stream crossing Glenn Highway at milepost 147.2, outlet of Snowshoe Lake, 
Aug. 14, 1950, moderate current, water somewhat colored, 17.0° C.; one young 
specimen, under a stone. 

Moose Creek, on Glenn Highway, milepost 186, near Alaska Road Commission 
camp, Aug. 14, 1950, fast, clear stream, 16.2° C; under stones, two mature speci- 
mens (holotype). 

Pippin Lake, on Richardson Highway, milepost 84.4, Aug. 15, 1950, shallow 
water near bank, 20° C. ; under stones, many specimens, two of them mature. 

Fresh-water lake near Umiat, collected by P. F. Scholander, Aug. 15, 1948; 
18 specimens, majority belonging to the plain form, 1 with distinct stripes, 5 
sexually mature (USNM 23678). 

Dendrocoelopsis alaskensis, new species 

Figure 24; Plate 6, Figure 4 

Description. — I had only limited material of this species and none 
of the animals was fully mature. The largest specimen measured 20 
mm. in length and 4 mm. in width. 

The anterior end is slightly lobed, with a convex frontal margin 



U. S. NATIONAL MUSEUM 



PROCEEDINGS. VOL., 103, PLATE 7 





Top: Beaver pond at milepost 62.6 of Richardson Highway, x\laska. Below the beaver 
dam, Polycelis borealis under stones. Bottom: Stream flowing along Richardson High- 
way, Alaska, at milepost 26.3. On the undersides of stones, Polycelis borealis. 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 103. PLATE 8 





Top: Stream crossing Sieese Highway, Alaska, at milepost 84.0. Habitat of Pkagocata 
nivea and Dendrocoelopsis alaskensis. Bottom: Lake at milepost 23.5 of Glenn High- 
way, Alaska. Under stones near bank, Dendrocoelopsis piriformis. 



FRESH-WATER TRICLADS OF ALASKA — ^KENK 179 

and a pair of rounded auricles protruding both anteriorly and later- 
ally. No distinct adhesive or grasping organ is developed. There is 
a gentle narrowing of the head (neck) behind the auricles. Behind 
the head, the width of the body gradually increases until the greatest 
width is reached. The posterior end is bluntly pointed. 

The body lacks pigment, being white except for the contents of 
the intestine, which may show through the body wall. 

There are two principal eyes. The distance between them amounts 
to about one-third the width of the head at the level of the eyes. 
The distance of each eye from the frontal margin is equal to, or 
slightly larger than, the distance from the lateral margin. Additional, 
supernumerary, eyes were seen in a few individuals at short distances 
either anterior or posterior to the principal eyes. 

The species bears a striking resemblance to a European species, 
Dendrocoelum nausicaae Schmidt of the Balkan Peninsula. It is 
generally associated with another triclad inhabiting the same streams 
in Alaska, Phagocata nivea, from which it may be distinguished by 
its larger size and by the position of the eyes. 

A distinct adhesive organ is not seen in living animals. In histologi- 
cal sections, a transverse band, pierced by numerous openings of 
eosinophilic glands, is found below the frontal margin of the head. 
The nature of the glands and the local differentiation of the epithelium 
correspond entirely to the structure of the submarginal adhesive zone 
which is seen in this species, as well as in other triclads, bordering the 
lateral margins of the body. The frontal adhesive area may, therefore, 
be interpreted as an extension of the lateral adhesive zone. It is 
somewhat wider than the lateral one and has no muscular differen- 
tiations such as are typical of true grasping organs. The continuity 
of the two zones is interrupted by a short gap on each side of the 
head below the auricle. 

The internal muscle zone of the pharynx consists of a layer of inter- 
mingled circular and longitudinal fibers. This character places the 
species in the family Dendrocoelidae. The anterior intestinal trunk 
bears a fairly large number, 21 to 24 pairs, of lateral branches, or 
diverticula. 

Of a total of eight individuals of this species at my disposal, five 
were young, two showed primordia of genital structures, and only one 
had the principal parts of the reproductive system developed, though 
not fully differentiated histologically. The description of the genital 
organs, therefore, must be given with certain reservations. 

The testes are predominantly ventral, situated mainly below the 
level of the intestinal diverticula, occasionally extending farther 
dorsally between the diverticula. No clearly recognizable testes were 
seen behind the level of the mouth; they may, however, appear there 



180 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

when the animal matures completely. The ovaries are situated 
behind the third or fourth lateral branch of the anterior iatestiual 
trunk. 

The genital pore (pg) leads into an undivided cavity, the genital 
atrium (a), which is narrow at the pore and expands anteriorly. The 
narrow posterior part receives the outlet of the copulatory bursa and, 
anteriorly to it, the opening of the common oviduct. 

The penis has a fauiy large, spherical bulb and a plump and short 
papilla. The bulb contains a cavity with irregular outline, the seminal 
vesicle (vs), into which the two vasa deferentia (vd) open separately. 
The cavity of the bulb continues into the broad papUla and opens at 
its tip. The bulb has the usual coat of muscle fibers arranged in 
concentric layers. The papilla, which projects only little into the 
genital atrium, has two muscle layers underlying the outer epithelium, 
a circular layer and a longitudinal one. No glandular structures are 
differentiated in my specimen. 

The two oviducts unite, without embracing the bursa duct, dorsally 
to the genital atrium. The common oviduct (ode) opens into the 
posterior, narrow part of the atrium from the dorsal side. 

The bursa (b) is, in my specimen, a rather small sac with a narrow 
lumen. The bursa duct (bd) runs posteriorly, above the penis, to a 
level behind the genital pore. There it turns abruptly toward the 
ventral side. Its terminal portion is considerably wider than the 
anterior part of the duct and opens from the posterodorsal side into 
the genital atrium close to the genital aperture. In full maturity, the 
widened section of the bursa duct probably represents a histologically 
distinct vagina. 

Taxonomic position. — The systematic relations of Dendrocoelopsis 
alaskensis are discussed together with those of the preceding species, 
D. piriformis. D. alaskensis is distinguished from all other species 
of the genus by the lack of a grasping organ. 

Holotype. On seven slides, USNM 22335, creek crossing Elliot 
Highway at milepost 31.0, July 24, 1950. 

Distribution and ecology. — Dendrocoelopsis alaskensis is an inhabitant 
of cool, fast streams of the White Mountains and usually shares its 
habitat with Phagocata nivea. It is a stenothermic and rheophUic 
species. 

Clear spring and creek on Steese Highway, milepost 82.5, at the Alaska Road 
Commission camp, July 21, 1950; one immature specimen, under a stone. 

Stream crossing Steese Highway at milepost 84.0 (see pi. 8), altitude 2,700 
feet, July 21, 1950; two immature specimens, under stones, near liver bait. 

Creek crossing Elliot Highway at milepost 31.0, July 24, 1950, water tempera- 
ture 3.2° C; under stones, five specimens, two of them with sexual structures 
Giolotype) . 



FRESH-WATER TRICLADS OF ALASKA — KENK 



181 




I I I 

a ode pg bd 

Figure 24. — Dendrocoelopsis alaskensis, diagram of the copulatory organs in longitudinal 
section, X 70. a, genital atrium; b, bursa; bd, bursa stalk; ode, common oviduct; 
pg, genital pore; ph, pharynx (extended through the mouth); vd, vas deferens; 
vs, seminal vesicle. 

Zoogeographic conclusions 

The occurrence in Alaska of four endemic species of triclads poses 
several interesting questions. It is known that fresh-water triclads 
are most abundantly represented in the northern Temperate Zone 
and that the number of species declines toward both the Arctic and 
the Tropical Zones. The paucity of the triclad fauna at high lati- 
tudes has frequently been attributed to the effects of glaciation in 
rather recent geologic time. During the glacial period, when huge 
ice masses covered great parts of the northern hemisphere, all fresh- 
water life over wide areas must have vanished. After the glaciers 
had receded, the areas were gradually repopulated by species entering 
them from adjacent territories. The pattern of distribution of 
freshwater triclads m Europe shows evidence of a definite succession 
of species which migrated into the previously glaciated areas in the 
postglacial period (cf. Thienemann's 1950 summary of the pertinent 
literature) . 

It is well known, however, that the greater part of Alaska was 
not covered with ice in the glacial period. Geologic evidence of 
glaciation has been found only in the Brooks Range, the Alaska 
Range (and areas south of it), and in small isolated spots in the 
Yukon and Kuskokwim River Valleys while the remaining surface 
of Alaska remained free of ice (cf. Geological Society of America, 



182 PROCEEDINGS OF THE NATIONAL MUSEUM vou i03 

1945). We are thus justified in assuming that the fresh-water life 
of Alaska was not completely destroyed even at the peaks of glacia- 
tion when practically all Canada and a considerable part of the 
United States were covered with ice caps. 

The Alaskan fresh-water triclads show no close relationships with 
the present North American triclad fauna inhabiting the midwestem 
plains and the eastern and southern areas of the continent. A 
comparison with the fauna of the Rocky Mountains is not possible 
at present, since the West of the United States and of Canada is 
almost unexplored with regard to triclads and to lower aquatic in- 
vertebrates in general. In any event it appears highly improbable 
that Alaska was populated by species migrating to it from the south 
or east. 

On the other hand, the relations of the Alaska triclads to the 
fauna of Eurasia are unmistakable. Though, according to our 
present knowledge, none of the Alaskan triclads is specifically identi- 
cal with any Asiatic form, two species, Phagocata nivea and Polycelis 
horealis, are very closely related to Asiatic species of the same genera. 

Figure 25 shows the geographic range of the genus Polycelis as it 
is known at present. The individual dots on the map represent 
either single records or groups of neighboring localities where species 
of the genus have been found. In interpreting the map it is to be 
kept in mind that not all geographical areas are equally well mvesti- 
gated with regard to the occurrence of lower invertebrates. Europe, 
Japan, and the eastern parts of the United States are comparatively 
well known while our knowledge of the Asiatic, Western American, 
and African triclads is still rather deficient. A study of the map 
suggests that Polycelis is primarily a Eurasian genus. It appears 
to have extended its range, in some earlier geological period, to the 
northern rim of Africa.* In a similar way it may have migrated into 
Alaska at the time of the cenozoic land bridges and may have pene- 
trated south along the Rocky Mountains. It is highly probable 
that Polycelis will be found, in the future, more widely distributed 
in the Rockies than present collection records indicate. 

The repeated emergence of a land bridge between Alaska and the 
eastern tip of Siberia is generally accepted by geologists and paleon- 
tologists. There is ample evidence of an exchange of faunal elements 
between the two continents (cf. Simpson, 1940 and 1947). The most 
recent corridor must have existed during the glacial stages of the 
Pleistocene. The volume of ice masses accumulated over vast areas 
of the northern hemisphere has been estimated conservatively at 
34 to 42 million cubic kilometers. The corresponding depletion of 

♦ The presence of Polycelis oculi-marginata (Palombi) in New Guinea is a zoogeographic enigma (cf. Beau- 
champ, 1947). 



FRESH-WATER TRICLADS OF ALASKA — KENK 



183 




fe 



184 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

the ocean must have lowered the sea level 50 to 90 meters below the 
present level (Daly, 1934, pp. 41-50). The depth of the eastern part 
of the Bering Sea (withm the peruneter passing through the eastern 
projection of Siberia, the Seward Peninsula of Alaska, and Nunivak 
and St. Lawrence Islands) averages less than 40 meters. Thus the 
ocean bottom between Alaska and Asia must have emerged with each 
glacial stage, forming a wide connection between the two continents. 
This process may have been enhanced by the uplifting of the area 
along the margin of the ice cap, brought about by plastic or elastic 
deformation of the earth crust under the weight of the ice masses. 

A land bridge between Alaska and Asia must have persisted for 
prolonged periods of time. Simultaneously, Alaska was discon- 
nected from other inhabitable areas of North America by broad ice 
fields. 

In view of the geological history of Alaska we may conclude that 
the present Alaskan fresh-water triclads are the remnants or successors 
of triclads that lived in Alaska in the glacial times and perhaps even 
in preglacial periods, and that they are, in all probability, of Asiatic 
origin. It may further be assumed that some Alaskan triclads have 
extended their range during the postglacial period and proceeded 
southeast along the Rocky Mountains.^ The occurrence of a species 
of Polycelis in Wyoming and South Dakota may have resulted from 
this migration. More light would be thrown on this question by a 
more thorough study of the triclad fauna of the Canadian and 
American Rockies. 

Literature cited 

Arndt, Waltheb 

1922. Untersuchungen an Bachtricladen. Ein Beitrag zur Kenntnis der 

Paludicolen Korsikas, Rumaniens und Sibiriens. Zeitschr. fur wiss. 

Zool., vol. 120, pp. 98-146, pi. 4. 
Bazikalova, a. fA. 

1947. Turbellaria-Triclada vostochnol Sibiri i Pribalkal'fa. Doklady Akad. 

Nauk SSSR, new ser., vol. 55, pp. 671-672. 
Beauchamp, Paul be 

1932. Biospeologica. LVI. Turbellaries, Hirudinees, Branchiobdellidfe. 

Deuxifeme s^rie. Arch. Zool. Exp^r. et Gen., vol. 73, pp. 113-380, 

pis. 6-8. 
1939. La syst^matique et I'^thologie des Fonticola (Turb. Triclades). Vgstnfk 

Ceskoslov. Zool. Spolecnosti v Praze, vol. 6-7, pp. 91-96. 
1947. Observations sur quelques Turbellaries du Mus6e royal d'Histoire 

naturelle de Belgique. Bull. Mus. Roy. Hist. Nat. Belgique, vol. 

23, No. 33, 11 pp. 

1949. Biospeologica. LXIX. Turbellaries (troisifeme s^rie). Arch. Zool. 

Exp^r. et G6n., vol. 86, Notes et Revue, pp. 50-65. 

1950. Nouvelles diagnoses de triclades obscuricoles. Bull. Soc. Zool. France, 

vol. 75, pp. 65-70. 

• I wish here to correct my assumption in a previous paper (Kent, 1943, p. 6) that all Canadian triclads 
had entered Canada from the south. 



FRESH-WATER TRICLADS OF ALASKA — ^KENK 185 

Bergendal, D. 

1890. Studien iiber nordische Turbellarien und Nemertinen. Ofv. Vet.- 
Akad. Forh., 1890, pp. 323-328. 
Daly, Reginald Aldworth 

1934. The changing world of the ice age. xxi + 271 pp., 6 pis. New Haven. 
Geological Society of America 

1945. Glacial map of North America. Special Papers, No. 60. 
Girard, Charles 

1894. Recherches sur les Planari^s et les N^mertines de I'Amerique du Nord. 
Ann. Sci. Nat., ser. 7 (Zool.), vol. 15, pp. 145-310, pis. 3-6. 
Hyman, Libbie H. 

1931. Studies on the morphology, taxonomy, and distribution of North 
American triclad Turbellaria. III. On Polycelis coronata (Girard). 
Trans. Amer. Micr. Soc, vol. 50, pp. 124-135. 

1934. Report on triclad Turbellaria from Indian Tibet. Mem. Connecticut 

Acad. Arts Sci., vol. 10, No. 2, pp. 5-12, pis. 1-2. 

1935. Studies on the morphology, taxonomy, and distribution of North 

American triclad Turbellaria. VI. A new dendrocoelid from 

Montana, Dendrocoelopsis vaginatus n. sp. Trans. Amer. Micr. 

Soc, vol. 54, pp. 338-345. 
1937. Studies on the morphology, taxonomy, and distribution of North 

American triclad Turbellaria. VII. The two species confused 

under the name Phagocata gracilis, the validity of the generic name 

Phagocata Leidy 1847, and its priority over Fonticola Komdrek 1926. 

Trans. Amer. Micr. Soc, vol. 56, pp. 298-310. 
Ijima, I., and Kaburaki, T. 

1916. Preliminary descriptions of some Japanese triclads. Annot. Zool. 

Japon., vol. 9, pp. 153-171. 
Kaburaki, Tokio 

1922. On some Japanese freshwater triclads; with a note on the parallelism 

in their distribution in Europe and Japan. Journ. Coll. Sci., Imp. 

Univ. Tokyo, vol. 44, No. 2, 71 pp., 1 pi. 
Kenk, Roman 

1930. Beitrage zum System der Probursalier (Tricladida paludicola). III. 

Zool. Anz., vol. 89, pp. 289-302. 

1936. Bemerkung zur Trikladenfauna Turkestans. Zool. Anz., vol. 115, 

pp. 76-80. 
1943. Notes on the planarian fauna of Canada. Canadian Field-Nat., 
vol 57, pp. 5-6. 
LivANov, N. A., and Zabusova, Z. I. 

1940. Planarii basselna Teletskogo ozera i novye dannye o nekotorykh 
drugikh sibirskikh vidakh (Paludicoles du Lac Teletzkoe et nouvelles 
donnees sur quelques formes sib^riennes) . Trudy Obshch. Estes- 
tvoisp. pri Kazan. Univ., vol. 56, pp. 83-159. 
MuTH, Anton 

1912. Beitrage zur Kenntnis der Gattung Sorocelis Grube. Mitt. Naturw. 
Ver. Steiermark, vol. 48, pp. 381-410. 
Palombi, a. 

1931. Turbellari della Nuova Guinea. In: R^sultats scientifiques du 

voyage aux Indes Orientales Neerlandaises de LL. AA. RR. le 
Prince et la Princesse Leopold de Belgique, vol. 2, No. 8, 14 pp., 
1 pi. (M6m. Mus. Roy. Hist. Nat. Belgique, hors s6ne). 



186 PROCEEDINGS OF THE NATIONAL MUSEUM vou 103 

Sabxjssowa, Z. (see also Zabusova, Z. I.) 

1929. Die Turbellarien der Kamtschatka-Halbinsel nach den Sammlungen 

der Rjabuschinsky-Expedition 1908-1909. Zool. Jahrb., Abt. 

Syst., vol. 57, pp. 497-536, pi. 4. 
Seidl, Heinrich H. 

1911. Beitrage zur Kenntnis centralasiatischer Tricladen. Zeitschr. ftir 

wiss. Zool., vol. 98, pp. 31-67, pis. 5-7. 
Simpson, George Gaylord 

1940. Mammals and land bridges. Journ. Washington Acad. Sci., vol. 

30, pp. 137-163. 
1947. Holarctic mammalian faunas and continental relationships during 

the Cenozoic. Bull. Geol. Soc. Amer., vol 58, pp. 613-687. 
Thienemann, August 

1950. Verbreitungsgeschichte der Siisswassertierwelt Europas; Versuch 

einer historischen Tiergeographie der europaischen Binnengewasser. 

XVI + 809 pp. Stuttgart. 
Zabusov", I. P. (Sabussow, H.; Zaboussoff, H.) 

1901. Zamietki po morfologii i sistematikie Triclada. II. O planarlfakh" 

Onezhskago ozera (Tricladenstudien. II. Zur Kenntniss der 

Tricladen des Onegasees). Protok. Zasfed. Obshch. Estestvoisp. 

Imp. Kazan. Univ., Prilozhenle No. 191, 18 pp. 
1911. Izsliedovaniia po morfologii i sistematikie planarlK ozera Balkala. 

I. Rod" Sorocelis Grube (Untersuchungen iiber die Morphologie 

und Systematik der Planarien aus dem Baikalsee. I. Die Gattung 

Sorocelis Grube). Trudy Obshch. Estestvoisp. Imp. Kazan. Univ., 

vol. 43, No. 4, 422 + 8 + 2 pp., 11 pis. 
1916. Rjabuschinskya schmidti n. g. n. sp., novyl vid" i rod" Tricladida 

paludicola iz" Kamchatki {Rjabuschinskya schmidti n. g. n. sp., 

espece et genre nouveau des Tricladida paludicola du Kamtchatka). 

Russ. Zool. Zhur., vol. 1916, pp. 273-286. 
Zabusova, Z. I. (Saboussoff, Z. ; see also Sabussowa, Z.) 

1936. Planarii Kamchatki (Les planaires paludicoles du Kamtchatka). 

Kazan. Gosud. Univ., Uchenye Zapiski, Zool., vol. 96, No. 7, pp. 

141-174. 



U. S. 60VERNHENT PRINTING OFFICEi Ifll 




PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



issued 



SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 



Vol. 103 Wajhington: 1953 No. 3323 

REVIEW OF THE INDO-PACIFIC ANEMONE FISHES, GENUS 
AMPHIPRION, WITH DESCRIPTIONS OF TWO NEW 
SPECIES. 



By Leonard P. Schultz 



During the summer of 1950 Dr. Arthur D. Welander, School o 
Fisheries, University of Washington, and I were engaged in studying 
reef fishes brought back from the Marshall Islands by the staff of the 
Applied Fisheries Laboratory of the University of Washington. 
Among this material was a specimen of anemone fish that we could 
not identify with any known species. After I returned to the U. S. 
National Museum, I reviewed the descriptions of all known species, 
compared that specimen with the numerous lots of Amphiprion in 
the National Museum, and found that it represented a new species. 
During March 1951, I studied the anemone fishes in the Museum 
of Comparative Zoology, Harvard University, and found another 
undescribed species from Mauritius. 

Descriptions and analyses of species referable to the genus Amphi- 
prion have been based on so few specimens, usually only one or two, 
that the problem of variability or constancy of the color pattern has 
been neglected. For most of the few hundred species among more 
than fifty fish families that I have studied in detail the basic color 
pattern has been observed to be fairly constant. It is of great value 
for the recognition of species, especially in the genus Amphiprion. 
Weber and de Beaufort (The fishes of the Indo-Australian Archipelago, 
vol. 8, pp. 330-348, 1940) recognized eight species, whereas we have 
distinguished fom-teen and there may be others recognizable when 
larger series are compared and additional characters studied. Fin 
ray counts were made on various species and these data are recorded 
in table 1. 

Except for original descriptions, no attempt was made to include 
all references to species referable to the genus Amphiprion. Whenever 

238537—53 187 



188 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

figures of species were found these have been mcluded in the synonymy. 
Most descriptions and records are not in sufficient detail to assign 
them to the correct species without examining the material on which 
the records were based. 

Genus Amphiprion Bloch and Schneider 

Amphiprion Bloch and Schneider, Systema ichthyologiae . . ., p. 200, 1801 

(genotype, Lutjanus ephippium Bloch). 
Prochilus (on Klein, 1775) Bleeker, Nat. Verh. Hollandsche Maatsch, Wet. 

Haarlem, ser. 3, vol. 2, No. 6, p. 20, 1877 (genotjrpe, Lutjanus ephippium 

Bloch). 
Actinicola Fowler, Journ. Acad. Nat. Sci, Philadelphia, ser. 2, vol. 12, p. 533, 

1904 (genotype, Lutjanus percula Lacep&de). 
Phalerebus Whitley, Mem. Queensland Mus., vol. 9, pt. 3, p. 216, 1929 (genotype, 

Prochilus akallopisos Bleeker). 

Key to the species of Amphiprion 

la. A white band (sometimes indistinct) along middorsal line from snout to 
dorsal origin or beyond along base of dorsal fin; total pectoral rays 17 to 
19; next to last dorsal spine about 1.3 to 1.5 in longest dorsal spine; no 
notable emargination in dorsal fin; scales on dorsal surface of head extend 
forward to a Une between front of orbits. 
2a. No vertical pale bars; dorsal rays about X,18 or 19; anal about 11,12 or 

13; pectoral 17 to 19 A. akalJopisos Bleeker 

26. A single vertical pale bar about 2 scales wide from nape to subopercle; 
dorsal rays about X,16; anal about 11,12 or 13; pectoral 17. 

A. perideraion Bleeker 
16. No white band along middorsal line. 

3a. Caudal fin with pale or dusky roundish center posteriorly edged with 
black; outer edges of caudal fin white; second pale or white bar from 
rear of spiny dorsal fin to anus with a triangular anterior projection 
under depressed pectoral fin; tips of pelvics black; a broad white 
color bar on head and on caudal peduncle always present. Dorsal 
fin deeply indented at rear of spiny part, next to last dorsal spine 
contained three times in longest dorsal spine; scales on dorsal surface 

of head do not extend forward of nape A. percula (Lacepede) 

36. Color not as in A. percula. 

4a. Central part of caudal fin black; outer edges of caudal fin broadly or 

narrowly edged with white; second pale bar, if present, without 

any projection anteriorly; pale bar on head present; next to last 

dorsal spine contained 1.2 to 2.0 times in longest dorsal spine; no 

notable emargination in dorsal fin. 

5o. Second pale bar represented dorsally on body by an ovate white 

area that continues anterodistally on soft dorsal fin; this white 

area does not extend below midlengthwise axis of body and 

never to anus; first pale bar 10 to 12 scales wide; anal fin black, 

except distally edged with white; spiny dorsal black; pelvics 

black; pectoral pale, except basally blackish; next to last dorsal 

spine contained about 1.2 to 1.5 in longest dorsal spine; no 

notable emargination in dorsal fin; scales on dorsal surface of 

head extend forward to a line between rear of orbits. 

A. laticlavius Cuvier and Valenciennes 



INDO-PACIFIC FISH GENUS AMPHIPRION — SCHULTZ 189 

56. Second pale bar continuous from dorsal part of body to region 
of anus. 
6a. Second pale bar broad, about 7 to 14 scales wide at level of 
lateral line (its width there contained 2 or less times in width 
of third black bar at level of lateral line) from whence it con- 
tinues posterodorsally on soft dorsal fin; pelvics dusky to 
blackish; spiny dorsal blackish; anal blackish, at least basally; 
scales on dorsal surface of head extend forward to a line 
between rear of orbits. 
7a. Caudal peduncle with broad white bar; pectoral fin pale dis- 
tally, basally dusky, 

A. chrysogaster Cuvier and Valenciennes 

76. Caudal peduncle black; no white bar; black coloration of 

posterior part of body continues on central part of caudal 

fin; pectoral fin dusky in basal third.A. poiymnus (Linnaeus) 

66. Second pale bar narrow, about 2 to 6 scales wide at level of 

lateral line, its width there contained 3.5 or more times in 

width of third black bar at level of lateral line; caudal fin black, 

narrowly edged with white; anal fin pale to blackish; scales 

on dorsal surface of head extend forward to over rear half of 

pupil; pelvics and pectorals pale; soft dorsal black, edged with 

white. 

8a. Second and third (peduncular) pale bars about 2 or 3 scales 

wide, their width at level of lateral line contained 7 to 

10 times in width of third black bar at level of lateral 

line; second pale bar not extending to distal edge of 

dorsal fin, ending on basal half of last dorsal spine; 

posterodorsal part of body blackish; anterior and ventral 

parts of body pale; spiny dorsal light dusky. 

A. tricinctus Schultz and Welander, new species 
86. Second and third (peduncular) pale bars 4 to 6 scales wide, 
their widths at level of lateral Line contained from 4 to 
6.5 times in width of third black bar at level of lateral 
line; second pale bar extends into distal half of dorsal but 
not quite to the edge of that fin; dorsal and posterior half 
of body blackish; anteroventral part of body pale; spiny 
dorsal dusky to blackish. 

A. mauritiensis Schultz, new species 

4a. Caudal fin plain pale or plain dusky, no black central blotch edged 

with white posteriorly; first pale bar on head present. 

9a. Second pale bar broad, about 7 to 10 scales wide at level of lateral 

line, and continuing to distal edge of spiny dorsal fin, thence 

posteriorly along distal edge of soft dorsal; caudal fin pale; 

caudal peduncle black, without the third pale bar, at least on 

adults; spiny dorsal, anal, and pelvic fins black; pectoral fin pale; 

next to last dorsal spine contained about 1.5 in longest dorsal 

spine; scales on dorsal surface of head extend forward to a line 

between rear of orbits A. sebae Bleeker 

96. Second pale bar, if present, not continuing to distal edge of dorsal 
fin; next to last dorsal spine contained 0.8 to 1.2 in longest dorsal 
spine; scales on dorsal surface of head extend forward to lines 
between rear edge of orbits to center of pupil. 
10a. First two pale bars typically present on adults. 



190 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

llo. First two pale bars broad, each about 5 to 7 scales wide at 
level of lateral line; third (peduncular) pale bar represented 
by a white bar, posteriorly edged with darkish, or the dark 
body color on anterior part of caudal peduncle may end 
abruptly, the white continuing on caudal fin; spiny dorsal 
black, dusky, or pale; soft dorsal and anal fins pale to black; 
pelvics pale to black, sometimes edged with black anteriorly; 
pectoral pale; next to last dorsal spine about 1.2 in longest 

dorsal spine A. xanthurus Cuvier and Valenciennes 

116. First two pale bars narrow, second usually narrower than first, 
the latter 3 to 4.5 scales wide and the first 3 to 6 scales wide, 
at level of lateral line; caudal peduncle blackish, without 
third pale bar on adults, sometimes pale bar is present on 
young; dark color of caudal peduncle gradually fading into 
pale color of caudal fin; spiny dorsal pale to dusky, soft 
dorsal pale; anal fin pale; pectoral pale; pelvics pale, except 
anterior edge black; second from last dorsal spine contained 
from 1.0 to 1.2 times in longest dorsal spine. 

A. bicinctus Riippell 
106. First pale bar usually present on head, about 4 to 6 scales wide at 
level of lateral line; second pale bar lacking, except sometimes 
on small young specimens. 
12a. Anal fin pale or partly pale; pelvics with some pale area. 
13a. Anal fin pale, except distally edged with fine black line; 
pelvics pale, except anterior edge black; breast pale; 
dorsal spines X, soft dorsal rays usually 17. 

A. ephippium (Bloch) 
136. Anal fin pale or distally pale with basal half (or less) 
dusky to blackish; pelvics pale distally and blackish 
ventrally, or inner rays pale and outer rays broadly 
blackish, except distal tips, which are pale; dorsal spines 
IX or X (more often IX than X), soft dorsal rays usually 

17 or 18 A. frenatus Brevoort 

126. Anal and pelvic fins black, no pale areas anywhere; dorsal 
spines X, soft dorsal rays usually 17. 

A. melanopus Bleeker 

Amphiprion akaJlopisos Bleeker 

Plate 9, Figure A 

Amphiprion akallopisos Bleeker, Nat. Tijdschr. Nederl. -Indie, vol. 4, p. 281, 

1853. 
Prochilus akallopisus Bleeker, Atlas ichthyologique . . . , vol. 9, pi. 400, fig. 3, 

1878. 
Phalerebus Whitley (new genus), Mem. Queensland Mus., vol. 9, pt. 3, p. 216, 

1929 (genotype, Prochilus akallopisus Bleeker = A. akallopisos Bleeker). 

This species is best recognized by the presence of a wide white 
band from snout along middorsal line of head, thence posteriorly 
along each side of base of dorsal fin, and ending on dorsal side of caudal 
peduncle; no white bars are present. Three specimens were studied: 
USNM 147130, from the Philippine Islands; USNM 82781, from the 
Fiji Islands; and MCZ 3308, from Sabang Bay. 



INDO-PACIFIC FISH GENUS AMPHIPRION — SCHULTZ 191 

Amphiprion perideraion Bleeker 

Plate 9, Figuke B 

Amphiprion perideraion Bleeker, Nat. Tijdschr. Nederl-Indie, vol. 9, p. 437, 

1855. — Montalban, Bur. Sci. Manila Monogr. 24, p. 16, pi. 4, fig. 1, 1928 

(Philippine Islands). 
Prochilus perideraion Bleeker, Atlas ichthyologique . . . , vol. 9, pi. 400, fig. 1, 

1878. 
1 Amphiprion rosenbergi Bleeker, Acta Soc. Sci. Indo-Neerl., vol. 6, p. 16, 1859; 

Atlas ichthyologique . . . , vol. 9, pi. 402, fig. 2, 1878. 

This species is best recognized by the narrow white band on the 
middorsal line of head, beginning between front of eyes and extending 
to dorsal origin, and a narrow white vertical bar on rear of head be- 
hind eye. Four specimens were studied: USNM 141032, 141033, 
and 147129, from the Marshall Islands and Borneo; and MCZ 33409, 
from Amboina. 

Amphiprion percula (LacepMe) 

Plate 9, Figure C 

Lutjanus percula LacepSde, Histoire naturelle des poissons, vol. 4, pp. 194, 239, 
240, 1802 (New Britain). 

Amphiprion tunicatus Cuvier and Valenciennes, Histoire naturelle des poissons, 
vol. 5, p. 399, pi. 132, fig. 2, 1830 (Vanicolo) .—Lesson, Voyage ... La 
Coquille . . . , Zoologie, vol. 2, pt. 1, p. 192, pi. 25, fig. 3, 1830 (Port Praslin, 
New Ireland; Doreh, New Guinea). 

Prochilus percula Bleeker, Atlas ichthyologique . . • , vol. 9, pi. 400, fig. 2, 1878. 

Anthias polymnus var. (non Linnaeus) Bloch, Naturgeschichte der auslandischen 
Fische, vol. 6, p. 103, pi. 316, fig. 3, 1792. 

Amphiprion percula Giinther, Journ. Mus. Godeffroy, vol. 15, Andrew Garrett's 
Fische der Siidsee, pt. 7, pi. 124, fig. A, 1881 (Samoan Islands). — Day, The 
fishes of India . . . , vol. 2, p. 379, pi. 80, fig. 4, 1878 (Andamans) .—Montal- 
ban, Bur. Sci. Manila Monogr. 24, p. 14, pi. 2, fig. 2, 1928 (Philippine Islands). 

Actinicola percula Aoyagi, Coral Fishes, Tokyo, pi. 37, fig. 2, 1943 (Kakure- 
Kumanomi); Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 4, No. 1, 
p. 175, pi. 9, fig. 2, 1941 (Japan). 

Amphiprion bicolor Castelnau, Proc. Zool. Acclim. Soc. Victoria, p. 92, 1873 
(Port Darwin). 

Actinicola bicolor (Castelnau) Whitley, Mem. Queensland Mus., vol. 9, pt. 3, 
p. 215, pi. 27, fig. 2, 1929 (Port Darwin). 

This is the most characteristically colored species in the genus 
Amphiprion, and is one of the commonest seen associated with the 
sea anemone Discosoma. The second white bar has a forward pro- 
jection under the depressed pectoral fin. No other species of Am- 
phiprion observed by me has the forward edge of the second dark 
bar with a deep concavity. The centers of the dark bars on the 
sides may be pale brown to blackish; all dark bars are black edged. 
I have studied 20 lots, totaling 35 specimens, from the Philippine, 
Solomon, Palawan, Schouten, and Morotai Islands. 



192 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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ephippium 

north Australia, near Darwin.. 

melanopus 

Marshall Islands 

Samoan and Paumoto Islands 

frenatus 

Philippine Islands 

Okinawa and Japan 

akallopisos 

perideraion 

chrysogaster 

bicinctus 

mauritiensis 

tricinctus 

xanthurus 

percxda 

sebae 



INDO-PACiriC FISH GENUS AMPHIPRION — SCHULTZ 193 

Amphiprion laticlavius Cuvier and Valenciennes 

Plate 9, Figure D 

Amphiprion laticlavius Cuvier and Valenciennes, Histoire naturelle des poissons, 

vol. 5, p. 394, pi. 132, fig. 1, 1830 (New Guinea). 
Amphiprion bifasciatus (non Bloch) Montalban, Bur. Sci. Manila Monogr. 24, 

p. 15, pi. 3, fig. 1, 1928 (Philippine Islands). 
Amphiprion bifasciatus annamensis Chevy, Travaux I'lnst. Oceanogr. Indochine, 

Mem. 4, pt. 1, poissons, p. 99, pi. 39, 1932 (Sud-Annam). 
Prochilus bifasciatus (non Bloch) Bleeker, Atlas ichthyologique . . . , vol. 9, pi. 400, 

fig. 4, 1878. 
Amphiprion polymnus (non Linnaeus) Aoyagi, Coral Fishes, Tokyo, pi. 36, fig. 2, 

1943 (Toaki-Kumanomi) ; Biogeographica, Trans. Biogeogr. Soc. Japan, 

vol. 4, No. 1, p. 173, pi. 12, fig. 4, 1941 (Japan). 
Amphiprion unimaculatus (non Meuschen) Okada and Ikeda, Biogeographica, 

Trans. Biogeogr. Soc. Japan, vol. 3, No. 2, p. 202, fig. 28, 1939 (Itoman, 

Okinawa) . 

This species differs from all others in regard to the second white 
bar, which is represented by an ovate white area on upper half of 
body and on soft dorsal fin, but without any extension on ventral part 
of body. Third white bar on caudal peduncle is lacldng. Caudal 
fin is basally and centrally blackish, with posterolateral edges white. 
Anal is submarginally blackish, with distal edge white. One specimen, 
USNM 147128, from the Philippines, was studied. 

Amphiprion chrysogaster Cuvier and Valenciennes 

Plate 9, Figure E 

Amphiprion chrysogaster Cuvier and Valenciennes, Histoire naturelle des poissons, 
vol. 5, p. 400, 1830 (lie de France) — Lesson, Voyage ... La Coquille, . . . , 
Zoologie, vol. 2, pt. 1, p. 191, pi. 28, fig. 3, 1830 (lie de France). 

Amphiprion percula (non Lacep&de) Okada and Ikeda, Biogeographica, Trans. 
Biogeogr. Soc. Japan, vol. 3, No. 2, p. 200, pi. 6, fig. 1, 1939 (Riu Kiu Islands), 

Amphiprion trifasciatus Cuvier and Valenciennes, Histoire naturelle des poissons, 
vol. 5, p. 695, 1830 (Moluccas). 

Amphiprion fusciventer Bennett, Proc. Comm. Zool, Soc. London, vol 1, p. 165, 
1831 (Mauritius). 

Prochilus bifasciatus (non Bloch) Bleeker, Atlas ichthyologique . . . , vol. 9, 
pi. 400, fig. 6, 1878. 

This species, with three broad white bars, has the central part 
of caudal fin black and outer edges white; second white bar is con- 
tinuous on distal part of soft dorsal fin. Seven specimens were 
studied: One each in USNM 61690 from Mauritius, 141034 from the 
Marshall Islands, 147127 from the Philippines; and four MCZ speci- 
mens, three from Zanzibar, Africa, and one, collected by Andrew 
Garrett, from Apiang, Kingsmill Islands. 



194 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

Amphiprion polymnus (Linnaeus) 

Plate 9, Figure J 

Perca ^olymna Linnaeus, Systema naturae, ed. 10, p. 291, 1758. 

Anthias bifasciatus Bloch, Naturgeschichte der auslandischen Fische, vol. 6, p. 

103, pi. 316, fig. 2, 1792. 
Prochitus bifasciatus Bleeker, Atlas ichthyologique . . . , vol. 9, pi. 400, fig. 5, 

1878. 
? Amphiprion ocellaris Cuvier and Valenciennes. Histoire naturelle des poissons, 

vol. 5, p. 399, 1830 (Sumatra). 
? Amphiprion melanurus Cuvier and Valenciennes, Histoire naturelle des poissons, 

vol. 5, p. 400, 1830 (Sumatra). 
Lutjanus jourdin LacepSde, Histoire naturelle des poissons, vol. 4, pp. 191, 235, 

1802 (Amboina). 
Coracinus seu Sciaena unimaculata Meuschen, Zoophylacium Gronovianum . . . , 

Pisces, No. 227, 1781 (based on Gronow, 1763). 
Coracinus vittatus Gray, Catalogue of fish collected and described by L. T. Gronow 

. . . , British Museum, p. 57, 1854 (based on Gronow's No. 227). 
Amphiprion intermedius Schlegel and Miiller, Verhandelingen over de Natuurlijke 

Geschiedenis der Nederlandsche Overzeesche Bezittingen . . . , Zoologie, 

p. 18, 1839-1841 (reference copied). 

Amphiprion polymnus (Linnaeus) has been confused almost since 
the day it was named. Weber and de Beaufort (Fishes of the Indo- 
AustraUan Archipelago, vol. 8, p. 344, 1940) discuss the confusion 
between polymnus and bicinctus, the former name having been used 
for the species currently called bicinctus. 

In A. polymnus the second white bar continues on to the distal 
part of the soft dorsal, but it lacks the third white bar on the caudal 
peduncle; also, the black coloration of the caudal peduncle continues 
on to the central part of the caudal fin, whereas in A. sebae the caudal 
fin is pale. The anal fin is black basally, with the distal third white. 



Explanation for Plate 9 

A, Amphiprion akallopisos Bleeker, photograph of a color drawing in the Philippine Albatross 
collection; B, A. perideraion Bleeker, photograph of a color drawing in the Philippine 
Albatross collection; C, A. percula (Lacepede), photograph of a color drawing in the 
Philippine Albatross collection; D, A. laticlavius Cuvier and Valenciennes, photograph 
of plate 36, figure 2, in Aoyagi, Coral Fishes, 1943 {= A. polymnus, non Linnaeus, AoyagI); 
E, A. chrysogaster Cuvier and Valenciennes, photograph of plate 6, figure 1, In Okada and 
Ikeda, Blogeographica, Trans. Blogeogr. Soc. Japan, vol. 3, No. 2, 1939 {^=A. percula, non 
Lacepede, Okada and Ikeda); F, Amphiprion sebae Bleeker, photograph of figure 27 in 
Okada and Ikeda^ Blogeographica, Trans. Blogeogr. Soc. Japan, vol. 3, No. 2, 1939; 
G, A. xanthurus Cuvier and Valenciennes, copy of figure 4 in Jordan and Dickerson, Proc. 
U. S. Nat. Mus., vol. 34, p. 611, 1908, of a specimen from Suva, Fiji; H, A. tricinctus, 
new species, holotype, USNM 152929, from Bikini Atoll, Amen Island, standard length 
75 mm.; I, A. mauritiensis, new species, holotype, MCZ 6093, from Mauritius, standard 
length 111.5 mm.; J, A. polymnus (Linnaeus), photograph of plate 316, figure 2, Bloch, 
Naturgeschichte der auslandischen Fische, vol. 6, 1792 {= Anthias bifasciatus 'Sioch). 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 103. PLATE 9 




Certain species of Amphiprion 
(For explanation, see facing page) 



U. S. NATIONAL MUSEUM 



PROCEEDINGS, VOL. 103. PLATE 10 



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INDO-PACIFIC FISH GENUS AMPHIPRION — SCHULTZ 195 

I have examined one specimen collected Jmie 16, 1948, by Dr. 
Robert R. Miller in "a submerged canoe on a sand bar in Little 
Lagoon, northeast end of Groote Eylandt, Gulf of Carpentaria, 
Australia." The color when alive was as follows: "The pale bars 
were coral pink, with narrow emerald borders; the dark bars were 
velvet-black." 

Amphiprion tricinctus Schultz and Welander, new species 

Plate 9, Figure H 

Amphiprion ephippium (non Bloch) var. chrysopterus (non Cuvier and Valen- 
ciennes) Giinther, Journ. Mus. Godeffroy, vol. 15, Andrew Garrett's Fische 
der Siidsee, pt. 7, pp. 224-225, pi. 122, fig. C, 1881 (Kingsmill Islands). 

Holotype. — USNM 152929, Bikini Atoll, Amen Island, lagoon, 
August 21, 1947, University of Washington, Staff of Applied Fisheries 
Laboratory, standard length 75 mm. 

Description. — Dorsal fin rays X,17; anal 11,14; pectoral ii,17,i to 
ii,16,ii; pelvics 1,5; branched caudal fin rays 8 + 7; vertical scale rows 
from upper edge of gill opening to base of caudal fin 54, scales between 
lateral line and base of first soft dorsal ray 5, and between lateral 
line and anal origin 20; pores in lateral line 36; predorsal scales 19 or 
20; gill rakers 5 + 1 + 13. 

Detailed measurements were made on the holotype and these data 
are expressed in thousandths of the standard length, 75 mm.: Greatest 
depth 560; length of head 272; snout 99; eye 95; least preorbital 35; 
length from snout tip to rear edge of maxillary 101 ; postorbital length 
of head 147; least width of interorbital space 100; least depth of 
caudal peduncle 160; length of caudal peduncle from base of last anal 
ray to midbase of caudal fin 192; length of longest ray of pectoral 287, 
pelvic 313, upper caudal fin 307, lower lobe of caudal fin 300, spiny 
dorsal 160; length of next to last dorsal spine 160; width of white 
part of first pale bar at level of lateral line 73, second 40, last (pe- 
duncular) 20. 

Depth of body 1.7, head 3.4, both in standard length. Snout 3.2; 
eye 3.1; least preorbital distance 8.0; upper jaw 2.8; postorbital part 
of head 2.0; least interorbital space 3.0; least depth of caudal peduncle 
1.8; length of pectoral fin 1.0, pelvic 0.9, second dorsal spine 23, 
upper caudal rays 0.9 to 1.0; all in length of the head. Least depth 
of caudal peduncle in its length 1.2. Angle of upper profile of head 
with lengthwise axis of body about 50° ; profile of head convex. 

Teeth in both jaws in a single row, nearly conical, a little com- 
pressed forward, pointed; interorbital space scaled forward to a line 
between middle of pupils ; 4 or 5 rows of scales on cheeks ; gill cover with 
a few scales; scales occur part way out on all median fins; preorbital 
with 3 spines; suborbital with 10 to 12 smaller spines. 



196 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Color in alcohol. — Background coloration of body from about fifth 
dorsal spine posteriorly and dorsally to midlengthwise axis of body 
blackish, the anteroventral part of body pale light brown becoming 
paler ventrally; spiny dorsal dark brown; soft dorsal black; caudal fin 
black, except edged with white posteriorly; pectoral and pelvic fins 
pale or very light tan ; anal pale or light tan, distally edged with a black 
line; first white bar begins a little in front of dorsal origin and just 
behind eye, ending on lower edge of subopercle; second white bar 
begins on last dorsal spine and base of first soft ray in lower third of 
fin, extends ventrally, meeting its fellow in narrow space between 
anal origin and anus; third white bar, about half width of second, or 
narrower than width of pupil, crosses caudal peduncle just in front of 
caudal fin base. 

Remarks. — This new species may be recognized by the narrowness 
of the three white bars, especially the second and third, and by the 
black caudal fin narrowly edged with white posteriorly. It is sep- 
arated from all other species of Amphriprion by the key. 

Named tricinctus in reference to the three white bars. 

Amphiprion mauritiensis Schultz, new species 

Plate 9, Figure I 

Holotype. — MCZ 6093, Mauritius, collected by Nicolas Pike, stand- 
ard length 111.5 mm. 

Paratypes. — Bearing same data as holotype: Out of MCZ 6093, 4 
specimens, 67 to 96 mm.; MCZ 5801, 2 specimens, 98 and 104.5 mm.; 
MCZ 5800, 1 specimen, 101.5 mm.; MCZ 5802, 2 specimens, 83.5 and 
114 mm. 

Description. — Dorsal rays X,17 or 18, one with XI, 16, usually 
X,17; anal 111,14, one with 111,13; pectoral rays 20 or 21; pel vies 
1,5; branched caudal fin rays 8+7; vertical scale rows from upper 
edge of gill opening to base of caudal fin about 55 to 58; scales between 
lateral line and base of first soft dorsal ray 5 ; between lateral line and 
anal origin 17 to 19; pores in lateral line 35 to 42; predorsal scales 
about 10 or 11; gill rakers 5 + 1 + 13. 

Detailed measurements were made on the holotype and two para- 
types, and these data are expressed in thousandths of the standard 
length, first for the holotype then, in parentheses, for the paratypes: 
Standard length in mm. 111.5 (87.5; 104.5). Greatest depth 485 
(490; 535); length of head 297 (286; 310); snout 90 (97; 96); eye 89 
(86; 86); least preorbital width 31 (38; 28); distance from snout tip 
to rear edge of maxillary 91 (114; 112); least width of interorbital 
space 74 (91; 81); least depth of caudal peduncle 144 (157; 151); 
length of caudal peduncle 166 (153; 148). Length of longest ray of 



INDO-PACIFIC FISH GENUS AMPHIPRION — SCHULTZ 197 

pectoral 279 (269; 280); pelvic 291 (257; 296); upper caudal lobe 270 
(280; 296); lower caudal lobe 265 (263; 277); spiny dorsal 135 (132; 
124). Length of next to last dorsal spine 99 (86; 86). Width of 
first white bar at level of lateral line 81 (80; 67); second 71 (74; 57); 
third (peduncular) 48 (63; 48). Width of second black bar at level 
of lateral line 323 (331; 344); third 300 (292; 344). 

Depth of body 1.8 to 2.1, head 3.2 to 3.5, both in standard length. 
Snout 2.9 to 3.2; eye 3.3 to 3.6; least preorbital 7.5 to 10.1; upper 
jaw 2.5 to 3.2; least interorbital 3.1 to 3.9; least depth of caudal 
peduncle 1.9 to 2.0, length of pectoral 1.1, pelvic 1.0 to 1.1, third 
dorsal spine 2.1 to 2.5; all in length of head. Least depth of caudal 
peduncle in its length 0.95 to 1.2. Width of second white bar at 
level of lateral line in width of thu'd black bar at level of lateral line 
3.5, thhd white bar 4.0 to 7.0. 

Profile of head convex. Teeth uniserial, nearly conical, a little 
compressed forward, pointed; interorbital space scaled forward to a 
line between rear of eyes; 4 to 6 rows of scales on cheek; gill cover 
with a few scales; scales occur part way out on all median fins; pre- 
orbital with 1 to 3 spines; suborbital with about a dozen spines. 

Color in alcohol. — Background coloration dark brownish or blackish, 
except that underside of head and ventrally below a line from pectorals 
to anus is white or pale; three white bars, one behind eye, second from 
origin of soft dorsal to anus, third on caudal peduncle; dorsal fin 
black, with distal edge of soft dorsal white; anal fin black or pale, if 
black, distal edge is white; pelvics pale; pectoral pale, except dusky 
basally on some specimens; central area of caudal fin blackish, edges 
white, with edge widest distally; gill membranes appear to be white. 

Remarks. — This new species is most closely related to A. tricinctus 
but differs in having wider white bars, as compared in the key. 
Named mauritiensis in reference to the locality where it was collected. 

Amphiprion sebae Bleeker 

Plate 9, Figure F 

Amphiprion sebae Bleeker, Nat. Tijdschr. Nederl.-Indie, vol. 4, p. 478, 1853; 
Atlas ichthyologique , . ., vol. 9, pi. 400, fig. 9, 1878. — Day, Fishes of 
India . . . , vol. 2, p. 378, pi. 80, fig. 3, 1878 (Andamans) .— Okada and 
Ikeda, Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 3, No. 2, p. 200, 
fig. 27, 1939 (Isigaki and Irimote Islands). 

This species, in which the second white color bar continues distally 
on the soft dorsal fin, is very much like A. polymnus, but differs in 
having a white bar on the caudal peduncle and a white caudal fin. 
Three specimens were studied: USNM 45169, from the Seychelles, 
and 133830, from the Paumotu Islands; and MCZ 33410, from Sumatra. 



198 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Amphiprion xanthurus Cuvier and Valenciennes 

Plate 9, Figure G 

Amphiprion xanthurus Cuvier and Valenciennes, Histoire naturelle des poissons, 

vol. 5, p. 402, 1830 (lie de France). 
Amphiprion clarckii Cuvier and Valenciennes, Histoire naturelle des poissons, 
ri vol. 9, p. 504, 1833 (emended spelling on Bennett). 
Anthias clarkii Bennett, A selection from . . . fishes found on the coast of Ceylon, 

London, ed. 2, p. 29, pi. 29, 1834 (Ceylon). 
Sparus milli Bory de Saint- Vincent, Dictionnaire classique d'histoire naturelle, 

vol. 17, p. 130, pi. 113, fig. 2, 1831 (China Sea). 
Prochilus polymnus (non Linnaeus) Bleeker, Atlas ichthyologique, vol. 9, pi. 400, 

figs. 7, 8, 1878. 
Anthias polymna (non Linnaeus) Bloch, Naturgeschichte der auslandischen 

Fische, vol. 9, p. 89, pi. 316, fig. 1, 1792. 
Amphiprion bicinctus (non Riippell) Aoyagi, Coral Fishes, Tokyo, pi. 37, fig. 1, 

1943 (Kumanomi). — Biogeographica, Trans. Biogeogr. Soc. Japan, vol, 4, 

No. 1, p. 169, pi. 9, fig. 1, 1941 (Japan). 
Amphiprion polymnus (non Linnaeus) Okada and Ikeda, Biogeographica, Trans. 

Biogeogr. Soc. Japan, vol. 3, No. 2, p. 204, fig. 30, 1939 (Riu Kiu Islands).— 

Montalban, Bur. Sci. Manila Monogr. 24, p. 10, pi. 1, fig. 1, 1928 (Philippine 

Islands) . 
Amphiprion chrysopterus Cuvier and Valenciennes, Histoire naturelle des poissons, 

vol. 5, p. 401, 1830 (no locality given). — Jordan and Dickerson, Proc. U. S. 

Nat. Mus., vol. 34, p. 612, fig.4, 1908 (Suva, Fiji). 
Amphiprion japonicus Temminck and Schlegel, Fauna Japonica . . . , Pisces, 

p. 66, 1843 (Japan). 
Amphiprion chrysargyrus Richardson, Rep. Meetings British Assoc. Adv. Sci., 

vol. 15 (1845), p. 254, 1846 (Seas of China and Japan). 
Amphiprion boholensis Cartier, Verh. phys. med. Wiirzburg, new ser., vol. 5, 

p. 96, 1874 (Bohol). 
Amphiprion melanostolus Richardson, Ann. Mag. Nat. Hist., vol. 9, p. 390, 1842 

(Depuch Island). 
7 Amphiprion de bojer Lienard, TreiziSme Rapp. Ann. Soc. Hist. Nat. Maurice, 

p. 68, 1843 (reference copied). 
Amphiprion snyderi Ishikawa, Proc. Nat. Hist. Tokyo Mus., vol. 1, No. 1, p. 11, 

pi. 5, 1904 (Bonin Island). 

I have examined 70 specimens (in 45 lots) of xanthurus, 22 of which 
are in the Museum of Comparative Zoology, Harvard University, 
and the others in the U. S. National Museum, and I find some variation 
in color pattern. Among these specimens 12 had black dorsal, anal, 
and pelvic fins; 3 had these three fins dusky; 10 had the dorsal and the 
anal black or dusky, with pelvics pale; 36 had the dorsal black and 
both anal and pelvics pale ; 9 had dorsal, anal, and pelvics pale, except 
in some specimens the pelvic fins were dark edged. Of these 70 
specimens 42 were from the Philippines, 4 were from Japan, 2 from 
the China coast, 18 from Zanzibar, 1 from the Gilbert Islands, and 
3 from Kingsmill Islands. 



INDO-PACIFIC FISH GENUS AMPHIPRION — SCHULTZ 199 

Amphiprion bicinctus Riippell 

Plate 10, Figure A 

Amphiprion bicinctus Riippell, Atlas zu der Reise im nordlichen Airika, p. 139, 

pi. 35, fig. 1, 1828 (Red Sea). 
Amphiprion papuensis Macleay, Proc. Linn. Soc. New South Wales, vol. 8, No. 

2, p. 271, 1883 (New Guinea). — Whitley, Mem. Queensland Mus., vol. 9, 

pt. 3, p. 210, pi. 27, fig. 1, 1929 (D'Entrecasteaux Group, New Guinea, on 

holotype) . 
Amphiprion arion De Vis, Proc. Linnean Soc. New South Wales, vol. 8, p. 450, 

1884 (South Seas). 

This species has the first two white bars but lacks the peduncular 
bar; the anterior edges of the pel vies are black and the caudal fin is 
pale. Six lots containing seven specimens were studied: USNM 
61679, from Suva, Fiji; USNM 141030 and 141031, from the Marshall 
Islands ; uncataloged specimens, one from the Paumotu and two with- 
out data; also one from Bikini in the University of Washington 
collection. 

Among these specimens, four adults, 82 to 99 mm. in standard length, 
have a pale anal fin, whereas two, 26 and 27 mm. long, have a black 
anal, and one, 25 mm. long, has a dusky anal fin. The caudal peduncle 
varies from pale dusky to black or brownish. The key gives the 
essential color pattern of this species. 

Amphiprion ephippium (Bloch) 

Plate 10, Figures B, C 

Luijanus ephippium Bloch, Naturgeschichte der auslandischen Fische, vol. 4, 

p. 121, 1790 (reference not seen); Ichthyologie, ou histoire naturelle, generate 

et particulifere des poissons . . . , vol. 7, p. 98, pi. 250, fig. 2, 1797 (East Indies). 
Amphiprion ephippium Day, Fishes of India . . . , vol. 2, p. 378, pi. 80, fig. 1, 

1878. 
Amphiprion monofasciatus Thiolliere in Montrouzier, Suite de la faune de I'lle 

de Woodlark ou Moiou, Ichthyologie, Ann. Sci., Physic. Nat. Agr. Indust., 

Lyon, vol. 8, p. 476, 1856 (Woodlark Island). 
Amphiprion tricolor Gtinther, Catalogue of the fishes in the British Museum, 

vol. 4, p. 8, 1862 (Port Essington; South Australia). 
Amphiprion riippeli Castelnau, Proc. Zool. Acclim. Soc. Victoria, vol. 2, p. 91, 

1873 (Port Darwin). 
Amphiprion frenatus (non Brevoort) Day, Fishes of India . . . , vol. 2, p. 378, 

pi. 80, fig. 2, 1878. 
Prochilos ephippium Bleeker, Atlas ichthyologique . . . , vol. 9, pi. 401, figs. 

1, 9, 1878. 
Amphiprion rubrocinctus Richardson, Ann. Mag. Nat. Hist., vol. 9, p. 391, 1842 

(Depuch Island; probably young). 

The adults of this species usually have a plain blackish body and 
head, with a single white color bar on the head, but on a few speci- 
mens, the largest in the series, this first white bar is lacking or nearly 



200 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

SO. The smallest specimens among 57 collected by Dr. Robert R. 
Miller from Woods Inlet west of Darwin, Australia, have a variable 
color pattern of two or three white bars. One specimen, 20 mm. in 
standard length, has only one white bar on the head; 9, from 20 to 
31 mm., have two white color bars, the first and second; 7, from 19.5 
to 28 mm., have the first and second white bars distinct and the third 
indistinct, sometimes represented by a white spot on dorsal edge of 
caudal peduncle; 10, from 17 to 25 mm., have all three white color 
bars distinct. I presume that the second and third white bars dis- 
appear with increase in size, since a 35-mm. specimen in this lot looks 
like all the larger specimens with only one white bar on the head. 

Amphiprion frenatus Brevoort 

Plate 10, Figure D 

Amphiprion frenatus Brevoort, U. S. Japan Exped. Nat. Hist., Washington, 
vol. 2, p. 263, pi. 6, fig. 4, 1856 (Lew Chew [Okinawa]). — Jordan and Snyder, 
Proc. U. S. Nat. Mus., vol. 24, p. 597, 1902.— Montalban, Bur. Sci. Manila 
Monogr. 24, p. 12, pi. 2, fig. 1, 1928 (Philippine Islands). — Aoyagi, Biogeo- 
graphica. Trans. Biogeogr. Soc. Japan, vol. 4, No. 1, p. 167, 1941 (Japan). — 
Okada and Ikeda, Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 3, No. 
2, p. 203, fig. 29, 1939 (Riu Kiu Islands). 

Prochilus polylepis Bleeker, Versl. Akad. Wet. Amsterdam, ser. 2, vol. 11, p. 135, 
1877; Atlas ichthyologique . . . , vol. 9, pi. 401, fig. 6, 1878. 

Amphiprion polymnus (non Linnaeus) Montalban, Bur. Sci. Manila Monogr. 
24, p. 10, pi. 1, fig. 1, 1928 (Philippines). 

I have studied 72 specimens from the Philippines, Okinawa, and 
Japan referable to this species. In alcohol two color phases are evi- 
dent, the usual one (53 specimens, ranging from 37 to 100 mm. in 
standard length) has a plain blackish body behind head, whereas 15, 
ranging from 39 to 60 mm., have 3 lengthwise pale bands on side of 
body, and 4 others are intermediate in regard to coloration. These 
pale bands at best are not very distinct. This species is character- 
ized by having about twice as many of the specimens with IX dorsal 
spines as with X; this latter figure is the usual number of dorsal 
spines in the other species of Amphiprion. 

Of 78 specimens in 54 lots studied, 76 were from the Philippine 
Islands and one each from Japan and Okinawa. The specimen from 
Okinawa (the type locality oi frenatus), USNM 71702, Naha, Okinawa, 
Albatross, 69 mm. in standard length, is herewith designated as neo- 
type of Amphiprion frenatus Brevoort since the type is not in existence. 



INDO-PACIFIC FISH GENUS AMPHIPRION — SCHULTZ 201 

Amphiprion melanopus Bleeker 

Plate 10, Fiqubb E 

Amphiprion melanopus Bleeker, Nat. Tijdschr. Nederl. Indie, vol, 3, p. 561, 1852. 
Prochilus melanopus Bleeker, Atlas ichthyologique . . . , vol. 9, pi. 401, fig. 7, 

1878. 
Prochilus macrostomus Bleeker, Atlas ichthyologique . . . , vol. 9, pi. 401, fig. 5, 

1878. 
Amphiprion ephippium (non Bloch) Giinther, Journ. Mus. Godeflfroy, vol. 15, 

pt. 7, pi. 122, fig. D (var. melanopus on p. 225), 1881. 
Amphiprion mccullochi Whitley, Mem. Queensland Mus., vol. 9, pt. 3, p. 213, 

1929 (Lord Howe Island). 
Amphiprion macrostoma (non Bleeker) Chevy, Travaux Inst. Oceanogr. Indo- 

Chine, Mem. 4, pt. 1, Poissons, p. 102, pi. 40, 1932 (Annam). 

This species is characterized by the single white bar on the head, 
plain black body, pale soft dorsal, caudal, and pectoral, and black 
pel vies and anal fins. 

Of 18 specimens in 7 lots studied, 14 were from the Marshall 
Islands, 3 from Apia, Samoa, and 1 from Paumotu Island. 



a. t. SOVEIMHEHT PKINTIH* OrFICEi 1*13 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 



issued i^§y\A, m'J^I h the 




SMITHSONIAN INSTITUTION 

U. S. NATIONAL MUSEUM 
Vol.103 Washington: 1954 No. 3324 

MARINE POLYCHAETE WORMS FROM POINT BARROW, 

ALASKA, WITH ADDITIONAL RECORDS FROM THE 

NORTH ATLANTIC AND NORTH PACIFIC 



By Marian H. Pettibone 



This report of the Arctic polychaetes found in the region of Point 
Barrow, Alaska, is based on material collected during 1948, 1949, and 
1950 by G. E. MacGinitie, of the Arctic Research Laboratory. Speci- 
mens were obtained from Eluitkak Pass, Elson Lagoon, near Point 
Barrow, were washed ashore at the Point Barrow base, and were 
dredged within 16 miles offshore at Point Barrow base in depths of 
1.7 to 123.5 fathoms on bottoms of mud, stones, gravel, rocks, in 
masses of worm tubes, and various combinations of these. Addi- 
tional specimens were collected from fish traps, from screen traps 
lowered through holes made in the ice, and from plankton hauls, 
some of which were made through holes in the ice. The latter collec- 
tions are of particular interest in that they include specimens showing 
sexual epitokous stages of some of the syllids. Considerable care 
was taken in going over the miscellaneous material and separating 
polychaetes, as evidenced by the large collection and the presence 
in it of many small specimens of the young of larger species as well 
as small species which are often overlooked. There are a great num- 
ber of small syllids, phyllodocids, hesionids, and terebeUids. Great 
care also was taken in the preservation of the specimens. Some 
color notes were included and some color photographs were taken, and 
these added considerably to the value of the collection. 

The material was worked on by the writer at the U. S. National 
Museum, where the collection is deposited. The facilities of the Mu- 
seum, including laboratory accommodations, the library, and the vast 
261112—54 1 203 



204 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

polychaete collections, which include much type material, were placed 
at the writer's disposal. Some previously unworked material was 
examined for comparative purposes, and the results of some of these 
related studies are included here. This material is chiefly from collec- 
tions made by the following: R. A. Bartlett in the Canadian Arctic, 
Greenland, and Labrador from 1927 to 1942; W. H. Dall in Arctic 
Alaska, the Bering Sea, and southeastern Alaska from 1871 to 1880; 
the Blue Dolphin expeditions to Labrador and Newfoundland from 
1949 to 1951 under the command of D. C. Nutt; the U. S. Fish Com- 
mission in dredgings off the east coast of North America, from which 
many specimens had been identified and recorded by A. E. Verrill; 
and by the writer in the Straits of Juan de Fuca, in Washington and 
Puget Sounds, Washington, chiefly during the summers from 1936 to 
1940, and from the region of Woods Hole, Massachusetts, in the 
summers of 1950 and 1951. For the last three collections mentioned, 
only summaries of the data are given for species common to the 
Point Barrow region. More complete data are to be published sepa- 
rately and the summaries are included here only to make the dis- 
tributional data of the Point Barrow species more complete. 

The number of polychaetes previously recorded from Arctic Alaska 
is smaU indeed. The collection obtained by the International Polar 
Expedition to Point Barrow from 1881 to 1883 included only 17 poly- 
chaetes (Murdoch, 1885, p. 152). This collection, which was depos- 
ited in the U. S. National Museum, was examined by the writer and 
is referred to in this report. A few additional records in scattered 
papers have added to the list of polychaetes of Arctic Alaska. 

For each of the species, a rather full but by no means complete 
synonymy is given, chiefly bringing together references to the original 
and additional descriptions and scattered distributional records. To 
facilitate identification, keys to the families, genera, and species are 
given, as well as synopses of the families and brief descriptions of the 
species with size ranges and color notes. An explanatory key to the 
lettering of the diagnostic features in the figures is given on page 2 10. 

This study was aided by a contract between the Office of Naval 
Research, Department of the Navy, and Johns Hopkins University 
(Project No. NR 162 911, Contract and Task Order No. N6onr 
243-16). 

The writer acknowledges her appreciation to Prof, and Mrs. G. E. 
MacGinitie for their cooperation and help on this project; to the au- 
thorities of the U. S. National Museum for allowing her to make use 
of the facilities of the Institution, especially to Dr. Waldo L. Schmitt 
and Dr. Fenner A. Chace, Jr., for their valuable aid, suggestions, and 
patience; and to Mr. and Mrs. Cyiil Berkeley, of Nanaimo, British 
Columbia, for their loan of specimens and their valuable suggestions. 



MARINE POLYCHAETE WORMS — PETTIBONE 205 

The distribution of the Point Barrow polychaetes is summarized 
in table 1. 

The collections include 3,270 specimens representing 88 species and 
26 families of Polychaeta. Some of the species were exceedingly 
common; others were represented only by single or few specimens. 
The Polynoidae, Syllidae, and Terebellidae are the most abundant 
both as to numbers of species (11 each, or 37 percent) and specimens 
(66 percent). The Phyllodocidae and Sabellidae have 7 species each. 
Eleven families are represented by a single species each. 

During 1948, because of the ice floes all summer, there was no 
heavy surf and practically nothing was washed up on the beach. 
During 1949, specimens were washed ashore on 17 different days. 
In 1950, specimens were collected from the beach on only one day. 
Altogether, 287 specimens of polychaetes were washed ashore, repre- 
senting 20 species and 11 families. Two species, Eunoe clarki (2 
specimens) and Travisia carnea (12 specimens), were obtained in this 
manner only. Four species were dredged occasionally but were 
washed ashore in much larger numbers, namely: Antinoe sarsi (58 
specimens), Alelaenis loveni (65 specimens), Arenicola glacialis (54 
specimens), and Brada villosa (24 specimens). Additional species 
were commonly obtained in the dredge but were washed ashore in 
small numbers. 

On three occasions in 1950 when 80-foot vertical plankton hauls 
were taken through holes in the ice, the sexual stages of three of the 
syllids were found — Avtolytits fallax, Syllis cornuta, and Syllis fasciata. 

A station 12.1 miles from shore, 123.5 fathoms, August 17, 1949, 
produced literally bushels of worm tubes. The tubes of the terebellid 
Pista maculata made up the greater part of the mass. 

Of the 88 Point Barrow species, 75 (85.2%) are common to the 
Arctic, Atlantic, and Pacific; 5 (5.7%) are common only to the Arctic 
and Pacific {Gattyana ciliata, Eusyllis magnijica, Pionosyllis compacta, 
Glycinde wireni, and Idanthyrsus armatus); 3 (3.4%) are common to 
the Arctic and Atlantic {Autolytus fallax, Eumida minuta, Travisia 
carnea); 4 (4.5%) are confined to the Arctic (Eunoe clarki, Nerinides 
sp., Arenicola glacialis, Ampharete vega); and 1 (1.1%) is bipolar, 
known only from the Arctic and Antarctic {Ammotrypane hreviata). 

The Point Barrow records help to complete the circumpolarity 
records for 58 circumpolar or almost circimipolar species, some of 
which previously had been known only from the Siberian and Canadian 
Arctic, Greenland, and Spitsbergen. The range has been extended 
for 28 species, 6 of which have been extended by combining Atlantic 
and Pacific specific names. 



206 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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SIGALIONIDAE (30 Specimens) 

Pholo6 minuta (Fabricius) 


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Eteone spetsbergensis Malmgren 

Phyllodoce groenlandica Oersted 

Eumlda minuta (Ditlevsen) 


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MARINE POLYCHAETE WORMS — PETTIBONE 



207 





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PROCEEDINGS OF THE NATIONAL MUSEUM 



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ScALiBREGMiDAE (17 Specimens) 

Scalibregma inflatum Rathke 

Ophelhdae (14 specimens) 

Ammotrypane breviata Ehlers 

Travisia carnea Verrill... 

CAPrTELLiDAE (26 Specimens) 

Capitella capitata (Fabricius).. 

Akenicolidae (67 specimens) 

Arenicola glacialis Murdoch.. 

Maldanidae (35 specimens) 

Praxillella praetermissa (Malmgren).. 

Maldane sarsi Malmgren 

Nicomache lumbricahs (Fabricius) 

Nicomache personata Johnson 

Petaloproctus tenuis (Thfiel) 

Sabellarudae (6 specimens) 

Idanthyrsus armatus Kinberg.. 

Sternaspidae (5 specimens) 

Stemaspis scutata (Ranzani)... 

Pectinarhdae (72 specimens) 

Pectinaria granulata (Linnfi) 

Pectinaria hyperborea (Malmgren) 

Ampharetidae (41 specimens) 

Ampharete vega (Wirfen) 

Ampharete acutifrons (Grube) 

Ampharete gogsi Malmgren 

Asabellides sibirica ( Wirgn) 



MARINE POLYCHAETE WORMS — PETTIBONE 



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210 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Explanation of symbols on figures 

Roman numerals indicate body segments. 



a, anus 

aBr, anal branchia 

aC, anal cirrus 

aCo, anal cone 

aCy, anal cylinder 

aK, anal knob 

aLi, anal ligule 

abd, abdominal region 

ac, aciculum (heavy spine buried in 
parapodial ramus) 

acH, acicular genital hook 

acL, acicular lobe 

ach, achaetous lobe 

ai, aileron of jaw 

an, antenna 

antLa, anterior or presetal lamella 

anta, antanal achaetous segment 

ba, barbule 

bl, blade of seta (terminal or end 
piece of composite or jointed seta) 

br, branchia 

brF, branchial filament 

brLo, branchial lobe 

brP, brood pouch 

buS, buccal segment 

cP, cephalic peak 

cCre, cephalic crest 

cPl, cephalic plate 

cR, cephalic ridge 

caPl, calcareous plate 

CO, collarette 

cph, cirrophore (base of a tentacular 
cirrus or a dorsal cirrus) 

cr, crotchet 

ere, crest 

dC, dorsal cirrus 

dL, dorsal lobe 

dN, dorsal notch 

dTu, dorsal tubercle (lobe correspond- 
ing to elytrophore on non-elytra- 
bearing segment) 

deR, denticled rim 

Dor, dorsal 

el, elytron or scale 

elph, elytrophore (lobe bearing ely- 
tron) 

ey, eye-spot 

fTu, facial tubercle 

frAn, frontal antenna 



frH, frontal horn 

h jaw 

lAn, lateral antenna 

IGr, lateral groove 

ILo, lateral lobe 

IN, lateral notch 

lOr, lateral sensory organ 

liPl, limbate plate 

loL, lower lip 

mAn, median antenna 

mLo, median lobe 

maxR, maxillary or distal ring 

mo, mouth 

nPa, nephridial papilla 

ne, neuropodium or ventral ramus 

neC, neuropodial cirrus 

neLi, neuropodial ligule 

neS, neuroseta (seta of neuropodium) 

neTo, neuropodial torus 

no, notopodium or dorsal ramus 

noLi, notopodial ligule 

noS, notoseta (seta of notopodium) 

noPi, notopodial pinnule 

noTo, notopodial torus 

nuEp, nuchal epaulette 

nuF, nuchal fold, lobe or collar 

nuG, nuchal groove 

nuH, nuchal hook 

nuO, nuchal organ 

nuPa, nuchal papilla 

oR, oral or basal ring 

oT, oral tentacles 

ocT, occipital tubercle 

op, operculum 

opPe, opercular peduncle 

p, palp 

pM, palmar membrane 

pa, papilla 

pal, paleae 

parF, parapodial flange 

para, paragnaths or horny denticles 

parath, parathoracic segment 

poL, postsetal or posterior lobe 

poLa, postsetal or posterior lamella 

pr, prostomium 

preL, presetal or anterior lobe 

pro, proboscis 

pyF, pygidial or anal funnel 

pyPl, pygidial plate 



MARINE POLYCHAETE WORMS — PETTIBONE 



211 



s, seta uL, upper lip 

scH, scaphal hook un, uncinus (very small and short, 

set, setigerous segment or setiger flattened setal hooks) 

setLo, setigerous lobe unPi, uncinigerous pinnule (flattened 

sh, shaft of seta (basal part of com- parapodial ramus bearing numerous 

posite or jointed seta) uncini or small setal hooks) 

spH, special hook unTo, uncinigerous torus (low, in- 

tC, tentacular cirrus flated parapodial ramus bearing 

tF, tentacular filament numerous uncini or small setal 

tM, tentacular membrane hooks) 

tS, tentacular segment or peristomium vC, ventral cirrus 

(anterior modified segments around vF, ventral faecal groove 

mouth) vL, ventral lobe 

th, thoracic region vPa, ventral papilla 

thM, thoracic membrane vSh, ventral shield 

to, tooth Ven, ventral 

tor, torus (low, inflated parapodial w, basal crenulate wing 

lobe) 

Order POLYCHAETA 
Key to the families of Polychaeta from Point Barrow 

1. Dorsal surface more or less completely covered by paired overlapping scales 

or elytra (fig. 26, a) 5 

Dorsal surface not covered by elytra 2 

2. Anterior end more or less concealed by feathery tentacles (fig. 38, a), chiti- 

nous golden setae forming an operculum (figs. 34, n; 35, d), long setae 
directed forward forming a cephahc cage, or long filamentous outgrowths 

(fig. 36, /) . With more or less permanent tubes 6 

Anterior end otherwise 3 

3. Body thickly papillated, grub-shaped 21 

Body smooth or finely granulated 4 

4. Prostomium with conspicuous antennae, with or without fleshy palps (figs. 

26-30), or prostomium or an anterior segment with pair of long, prehensile, 

tentacular palps (fig. 32, i) 13 

Prostomium without palps and without conspicuous antennae (fig. 32, a) or 
with 2 pairs minute antennae (figs. 30, j; 31, o, e) 18 

5. Elytra on all segments of posterior region (at least from segment 29 on). 

Dorsal cirri lacking Sigalionidae (p. 228) 

Elytra not on all segments of posterior region. With paired dorsal cirri on 
segments without elytra (fig. 26, a) Polynoidae (p. 213) 

6. Anterior end with branchial plume (fig. 38, a) 7 

Anterior end without branchial plume 8 

7. Branchial filaments without operculum. Without thoracic membrane. 

Tube gelatinous, membranous, or horny, covered or not with mud, sand, 

gravel, and debris Sabellidae (p. 332) 

Usually 1 or 2 opercula on modified branchial filaments (fig. 39, u) . Usually 
with thoracic membrane (fig. 39, u). Tube calcareous.. Serpulidae (p. 342) 

8. Anterior end with chitinous, golden setae forming an operculum (figs. 34, 

n; 35, d). In free, conical, sandy tubes or concreted sandy tubes attached 

to rocks, shells, etc 12 

Anterior end without opercular setae. Tube otherwise 9 



212 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

9. Setae of anterior segments directed anteriorly, forming a cephalic cage. 
Body covered with papillae and often with adherent sand grains; may be 

encased in thick mucoid sheath Flabelligeridae (p. 288) 

Anterior end without cephalic cage but with long, filamentous outgrowths 
(may have 2 groups of paleal setae directed forward; fig. 36, a, e, /). Body 
without papillae 10 

10. Anterior end with long, filamentous outgrowths and with similar outgrowths 

continued along sides of body on few to many segments (fig., 33, a). Seg- 
ments similar, body not divided into 2 distinct regions. 

Cirratulidae (p. 285) 

Long, filamentous outgrowths concentrated on anterior region only (figs. 36, 

37). Body with 2 distinct regions 11 

11. With numerous filiform oral tentacles retractile in mouth (fig. 36, e). Ante- 

rior region with or without 2 groups of paleal setae directed forward (fig. 

36, a, b) Ampharetidae (p. 314) 

With numerous, grooved, oral tentacles not retractile in mouth (fig. 36, /, g, 
i) . Anterior region without paleal setae Terebellidae (p. 318) 

12. Opercular setae forming 1-3 concentric rows (fig. 34, n-q). Tube of con- 

creted sand or fine gravel, attached Sabellariidae (p. 307) 

Opercular setae forming 2 bundles in horizontal rows, directed obliquely 
anteriorly (fig. 35, c, d). Tube free, conical, rigid, open at both ends, 
formed of single layer of sand grains Pectinariidae (p. 310) 

13. Prostomium with 2-5 conspicuous antennae. With 1-8 pairs tentacular cirri 

(figs. 27-29) 14 

Prostomium without conspicuous antennae. Without tentacular cirri. With 
pair of tentaculiform, longitudinally-grooved palps (deciduous, may be 
missing; figs. 32, i; 33, d) 17 

14. Dorsal cirri flattened, foliaceous, sometimes overlapping, or more or less 

globular (fig. 27). Prostomium with 2 eyes, 4-5 antennae, without 

palps Phyllodocidae (p. 231) 

Dorsal cirri otherwise. Prostomium ^vith 4-6 eyes, 2-3 antennae, 2 palps 
(may be reduced and fused in Syllidae) 15 

15. With parapodial ligules, rounded, conical or straplike lobes (fig. 30, h, c). 

Dorsal cirri short, cirriform Nereidae (p. 263) 

Without parapodial ligules. Dorsal cirri filiform, may be more or less articu- 
lated (or small and subulate in some Syllidae) 16 

16. Palps biarticulate (fig. 28, a) Hesionidae (p. 239) 

Palps not biarticulate (maj^ be reduced and fused; figs. 28, c; 29, a). 

Syllidae (p. 240) 

17. Parapodia with postsetal lamellae more or less developed (fig. 32, g). Branch- 

iae usuall.y paired, dorsal, ligulate or cirriform (rarely pinnate, rarely 

lacking) Spionidae (p. 280) 

Parapodia with lobes scarcely projecting, without postsetal lamellae. Branch- 
iae simple, filamentous, contractile, inserted above notopodia on few to 
many segments (fig. 33, d) Cirratulidae (p. 285) 

18. Prostomium with 2 pairs minute antennae (figs. 30, j; 31, o) 19 

Prostomium without antennae 22 

19. Prostomium subquadrate, with small antennae on anterior part (fig. 30, j). 

With anterior and posterior parapodial lamellae more or less developed 

(fig. 30, k-n) Nephtyidae (p. 266) 

Prostomium conical, pointed, annulated, with tiny terminal antennae (fig. 
31, o, e). Without parapodial lamellae 20 



MARINE POLYCHAETE WORMS — PETTIBONE 213 

20. Segments bi- or triannulate. Parapodia of single form throughout length of 

body, either only uniramous or only biramous Glyceridae (p. 272) 

Segments uniannulate. Body divided into 2 regions — anterior uniramous, 
posterior biramous (fig. 31,/, g) Goniadidae (p. 274) 

21. Posterior part of body with 2 horny ventral plates with radiating bundles of 

setae (fig. 35, 6) Sternaspidae (p. 309) 

Posterior part of body without horny plates Flabelligeridae (p. 288) 

22. Some of segments considerably elongated, much longer than wide. Anal 

segment with limbate plate, spatulate, or funnellike (fig. 34, d, h, rn). 
Prostomium hoodlike (fig. 34, i, I) or with limbate plate (fig. 34, b-c, g). 

Maldanidae (p. 302) 
Segments not much longer than wide 23 

23. Proboscis provided with dark, chitinous jaw pieces (fig. 31, i). Body smooth, 

elongate, cylindrical, resembling an earthworm. Parapodia weakly devel- 
oped simple lobes Lumbrineridae (p. 275) 

Proboscis without chitinous jaws. Form variable 24 

24. Body divided into 2 weakly to sharply separable regions, short anterior region, 

more or less flattened, with cushionlike neuropodia with several rows of 
setae, and long cylindrical posterior region with parapodial lobes projecting 

dorsally (fig. 32, c) Orbiniidae (p. 278) 

Parapodial lobes of median and posterior regions not projecting dorsally.. 25 

25. Prostomium conical 26 

Prostomium otherwise 27 

26. Body short, grub-shaped or lancet-shaped. Parapodia with simple capillary 

setae only Opheliidae (p. 295) 

Body elongate, slender, fragile. Parapodia with capillary setae and, in long 
posterior region, with rows of hooded hooks on somewhat inflated tori 
(fig. 33, t) Capitellidae (p. 298) 

27. Arborescent branchiae limited to anterior few segments. With capillary and 

forked parapodial setae Scalibregmidae (p. 293) 

Arborescent branchiae on long middle region of body. With capillary setae 
and sigmoid hooks embedded in elongated tori Arenicolidae (p. 300) 

Family Polynoidae 

Prostomium bilobed, with two pairs of eyes in trapezoidal arrange- 
ment, three dorsal antennae, a pair of subulate ventral palps. First 
or tentacular segment with setae reduced in number (0-3), with two 
pairs of tentacular cirri (fig. 26, a). Parapodia biramous, with setae 
simple (not compound), paired dorsal scales or elytra on certain seg- 
ments and paired dorsal cirri on non-elytra-bearing segments, paired 
subulate ventral cirri (fig. 26, a, d). Pygidium or anal segment with 
pair of anal cirri. Paired segmental or nephridial papillae near 
ventral bases of parapodia. Muscular proboscis eversible, with circle 
of soft marginal papillae and two pairs of interlocking, chitinous jaws. 

Represented by 7 genera and 11 species. All the genera con- 
sidered here have 15 pairs of elytra on segments 2, 4, 5, 7, 9, 11, 13, 
15, 17, 19, 21, 23, 26, 29, 32; lateral antennae inserted ventral to 
median antenna on prostomium (fig. 26, a). 



214 PROCEEDINGS OF THE NATIONAL MUSEUM vol,, los 

Key to the genera of Polynoidae from Point Barrow 

1. Notosetae stouter than or at least as stout as neurosetae, with tips blunt to 

pointed, not capillary 2 

Notosetae not as stout as neurosetae 5 

2. Notosetae large, smooth or with only faint transverse striations, few in number 

(5-9) Melaenis (p. 214) 

Notosetae with transverse spinous rows, more numerous (more than 10) 3 

3. Neurosetae long, slender, at least some end in capillary tips.Antinoe (p. 215) 
Neurosetae stouter, with tips straight or slightly hooked, not capillary 4 

4. All neurosetae with entire bare tips Eunoe (p. 216) 

Some of neurosetae bidentate, with secondary tooth well developed or rudi- 
mentary (fig. 26, e) Harmothoe (p. 220) 

5. Some notosetae with capillary tips. Segments less than 40 (35-38), thus, 

posterior end of body covered with elytra 6 

None of notosetae with capillary tips. Segments more than 40 (45-65), thus, 
a rather long posterior end without elytra. Elytra smooth, without 
tubercles Enipo (p. 225) 

6. Neurosetae of 2 kinds, upper few end in slender tips, rest end in bifid tips. 

Elytra smooth except for few microtubercles on anterior curved part. 

Arcteobia (p. 225) 

Neurosetae all similar, with entire tips. Elytra with microtubercles and 

sometimes also with macrotubercles Gattyana (p. 622) 

Genus Melaenis Malmgren, 1865 
Melaenis loveni Malmgren, 1865 

Melaenis lovini Malmgren, 1865, p. 78, pi. 10, fig. 10.— Th^el, 1879, p. 22.— 
Wir^n, 1883, p. 391, pi. 28, fig. 4; pi. 29, fig. 3.— Murdoch, 1885, p. 152.— 
Moore, 1908, p. 337.— Fauvel, 1914, p. 49.— Augener, 1928, p. 695.— 
Ditlevsen, 1937, p. 16.— Gorbunov, 1946, p. 38.— Wesenberg-Lund, 1950a, 
p. 9; 1950b, p. 29. 

Melaenis lov6ni var, gigantea Wir4n, 1883, p. 391, pi. 28, fig. 3; pi. 29, fig. 4. — 
Murdoch, 1885, p. 153. 

Description. — ^Ijength 25-76 mm., width including setae 9-22 mm- 
(up to 110 mm. long — Ditlevsen, 1937). Segments 39-41. Body 
elongated, thick, tapering both anteriorly and posteriorly. Pro- 
stomium without cephalic peaks, antennae smooth. Dorsal cirri 
with fine, scattered, clavate papillae. Elytra smooth, soft, trans- 
lucent to opaque, not covering middorsum and posterior few segments. 
Notosetae much stouter than neurosetae, few in number (5-9), 
light to dark amber-colored, smooth or with only very faint transverse 
striations. Neurosetae of two kinds: Numerous, long, slender, 
finely denticled, with capillary tips; few, shorter, with subequal, 
diverging, forked, blunt tips (one part of fork may be longer than the 
other). Color: In life and in alcohol: Wide greenish brown to 
bluish gray transverse stripes middorsally, green and white striped on 
dorsal tubercles, cirrophores and elytrophores; without color ventrally; 
elytra light to dark greenish in a wide, circular band, a circular area 



MARINE POLYCHAETE WORMS — PETTIBONE 215 

and lateral border without color; one specimen (collected October 4, 
1949) was noted as peach-colored from segment 10 posteriorly, the 
color showing mainly on the ventral side and extending into the 
parapodia (probably developing eggs). 

New records. — Arctic Alaska: Point Barrow base, washed ashore, 
1949 (July 30; August 21; September 22, 28, 30; October 4, 5, 16, 
17; 65 specimens); off Point Barrow base, along shore, 1.7 fms., on 
bottom of gravel with mud (2 specimens). East Coast North 
America: Off Labrador, 45 fms., Blue Dolphin Expedition, 1949. 

Distribution.— Widely distributed m the Arctic: Siberian and 
Alaskan Ai'ctic, Davis Strait, Greenland, Spitsbergen, Franz Josef 
Land, Kara Sea. Also Bering Sea; off Labrador. In 1.7-111 
fathoms. 

Genus Antinoe Kinberg, 1855 (sensu Malmgren, 1865) 
Antinoe sarsi Malmgren, 1865 

Antinoe sarsi Kinberg, 1862 (MS.), p. 468 (fide Malmgren, 1865).— Malmgren, 

1865, p. 75, pi. 9, fig. 6 (part) .—Mcintosh, 1900, p. 365, figs.— Chamberlin, 

1920, p. 8. — Hartman, 1944a, p. 334. 
Polynoe sarsi Theel, 1879, p. 16, pi. 1, fig. 8.— Wir6n, 1883, p. 390.— Murdoch, 

1885, p. 152. 
Harmothoe sarsi Eliason, 1920, p. 20.— Annenkova, 1931, p. 203.— Friedrich, 

1939, p. 122.— Thorsou, 1946, p. 48.— Wesenberg-Lund, 1950a, p. 7; 1950b, 

p. 23; 1951, p. 17. 
Harmothoe (Antinoella) sarsi Augener, 1928, p. 687. 
Antinoella sarsi Annenkova, 1937, p. 153; 1938, p. 137.— Gorbunov, 1946, p. 38.— 

Zatsepin, 1948, p. 107, pi. 28, fig. 8. 
Description.— Length. 21-68 mm., width mcluding setae 11-27 mm. 
Segments 37 or 38. Fragments easily, posterior segments frequently 
regenerating. Body elongate, flattened, tapering shghtly anteriorly 
and posteriorly. Prostomium with cephalic peaks distinct or poorly 
developed; anterior pan- of eyes larger than posterior, slightly anterior 
to greatest width of prostomium. Antennae with short, clavate 
papillae. Dorsal cirri extending beyond setae, long, slender, tapering 
gradually, with short, clavate papillae. May be a single anal ckrus, 
extremely long, thick at base, tapering gradually; may be one long 
and one short anal cirrus. Elytra large, thin, soft, smooth, with 
scattered microtubercles and short, delicate, clavate papillae. Lose 
elytra readily; often show regenerating elytra, thus appearing variable 
in size. Notopodia and neuropodia extending into conspicuous, 
digitiform, acicular lobes. Setae yellowish or golden. Notosetae 
much thicker than neurosetae, large, clear basally; more distally, 
finely spinous, pointed. Neurosetae of two kinds, with capillary tips 
and with slender, relatively obtuse, shghtly curved tips. Color: 
In life and preserved: Grayish gi-een or light brown middorsally, 



216 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i03 

including elytrophores and dorsal tubercles; ventral surface without 
color or dusky midventrally; elytra gray, translucent to opaque, 
with a darker brownish-gray medial crescent-shaped area, with a 
colorless circular area medial to and a darker area posterior to the 
place of attachment. 

Remarks. — A. sarsi differs from A. badia (Theel), with which it has 
sometimes been confused, by having the neurosetae of two lands, 
with capillary tips and with slender, relatively obtuse tips. In A. 
badia, all the neurosetae have capillary tips. 

Parasites. — One of the 63 specimens had the parasitic copepod 
Herpyllobius arcticus Steenstrup and Liitken attached to the prosto- 
mium (identified by P. L. Illg). 

New records. — Arctic Alaska: Point Barrow base, washed ashore 
1949 (August 21, 24; September 1, 12, 20, 22, 24, 26; October 5, 16; 
58 specimens) ; off Point Barrow base, up to 5 miles from shore, 5 to 
30.7 fms., on bottoms of mud or stones, and in screen trap through 
hole in ice, 6 fms., April 11, 1949 (3 stations, 5 specimens); near 
Point Belcher, Icy Cape, off Point Barrow, Dall. Bering Sea: 62°15' 
N., 167°48' W., 20 fms., G. M. Stoney, 1884. East Coast North 
America: Off Labrador, 6-30 fms., Blue Dolphin Expeditions, 
1950, 1951. 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Jan Mayen, Spitsbergen, Norway, Franz 
Josef Land, Barents Sea, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes to Great Britain, the Baltic; Labrador to Maine; Bering Sea; 
north Japan Sea. In 3-1,215 fathoms. 

Genus Eunoe Malmgren, 1865 
Key to the species of Eunoe from Point Barrow 

1. Elytra with scattered, bluntly conical microtubercles only, lacking fringe of 

papillae. Eyes small, anterior pair anterolateral on prostomium (scarcely 
visible dorsally) E. clarki 

Elytra with macrotubercles in addition to microtubercles, with lateral fringe 
of papillae. Eyes larger, anterior pair anterodorsal (visible dorsally) 2 

2. Body broad, oval, flattened. Extra rounded lobes on inner sides of elytro- 

phores and dorsal tubercles (lobes corresponding to elytrophores on non- 
elytra-bearing segments, fig. 26, c). Nuchal fold posterior to prostomium 
not prominent. Setae yellow. Elytra with microtubercles rather low, 
flattened, semiglobose, some bifid; macrotubercles confined essentially to 
single row near external border, nodular, with roughened tips. Antennae 

with short papillae E. nodosa 

Body more elongate, not so flattened. Without extra rounded lobes on inner 
sides of elytrophores and dorsal tubercles. Nuchal fold prominent. Setae 
dark amber-colored. Elytra with microtubercles one- to many-pronged; 
macrotubercles variable in number, size, and position, branched. Antennae 
with longer papillae E. oerstedi 



MARINE POLYCHAETE WORMS — PETTIBONE 217 

Eunoe clarki Pettibone, 1951 

Eunoe clarki Pettibone, 1951, p. 44, fig. 1. 

Description. — ^Length 36-38 mm., width including setae 12 mm. 
Segments 40 or 41. Body linear-oblong, tapered slightly anteriorly 
and posteriorly. Prostomium without cephalic peaks ; anterior pair of 
eyes anterolateral. Antennae and dorsal cirri with short, clavate 
papillae. Elytra nearly cover dorsum, with conical micro tubercles, 
without fringe of papillae. Color: In alcohol: Middorsum trans- 
versely banded grayish green; elytra greenish gray, with a darker 
mottled pattern on most of the exposed parts, with a darker spot 
medial to a lighter area over the elytophore; dorsal cirri with pig- 
mented bands basally and subterminally. 

New record. — Arctic Alaska: Point Barrow base, washed ashore, 
October 17, 1949 (2 specimens). 

Distribution. — Arctic Alaska. 

Eunoe nodosa (Sars, 1860) 

Figure 26, c 

Polynoe nodosa Sars, 1860, p. 58. 

Eunoe nodosa Malmgren, 1865, p. 64, pi. 8, fig. 4. — Fauvel, 1923, p. 51, fig. 18, 

a-e (part).— Annenkova, 1937, p. 149; 1938, p. 134.— Treadwell, 1937, p. 

27.— Friedrich, 1939, p. 122.— Berkeley and Berkeley, 1942, p. 187; 1943, 

p. 129.— Hartman, 1944a, pp. 334, 337.— Gorbunov, 1946, p. 38.— Zatsepin, 

1948, p. 107, pi. 28, fig. 7. 
Eunoa nodosa Malmgren, 1867, p. 6. — Webster and Benedict, 1884, p. 700; 1887, 

p. 708.— Mcintosh, 1900, p. 291, figs. 
Polynoe scabra Th^el, 1879, p. 7.— Wir6n, 1883, p. 387, pi. 28, fig. 2; pi. 29, fig. 

1 (part; not pi. 28, fig. 1; not Aphrodita scabra Fabricius, 1780). 
Polynoe islandica Hansen, 1882, pp. 17, 24, pi. 1, figs. 15-21.— Murdoch, 1884, 

p. 152. 
Polynoe arctica Hansen, 1882, pp. 21, 27, pi. 3, figs. 1-5. 
Polynoe spinulosa Hansen, 1882, p. 28, pi. 1, figs. 6-10. 
Polynoe foraminifera Hansen, 1882, pp. 21, 23, 29, pi. 1, figs. 11-14. 
Harmothoe nodosa Moore, 1902, p. 271. — Ditlevsen, 1917, p. 6, pi. 3, fig. 10 (part; 

not pi. 2, fig. 1).— Augener, 1928, p. 684 (part) .— Wesenberg-Lund, 1950a, 

p. 6 (part); 1950b, p. 17 (part); 1951, p. 10 (part). 
Eunoe depressa Hartman, 1948, p. 14 (part; not E. depressa Moore, 1905). 

Description.- — Length 13-75 mm., width including setae 7-35 mm. 
Segments 36 or 37. Prostomium with cephalic peaks short and blunt 
or lacking; anterior pair of eyes anterolateral, visible dorsally. An- 
tennae with short papillae. Dorsal cirri with long papillae. Body 
flattened ventrally, strongly arched dorsally, especially in anterior 
part. With characteristic bulbous structures on inner sides of bases 
of elytrophores and dorsal tubercles. Segmental papUlae quite elon- 
gate, cylindrical, directed dorsally between the parapodia. Elytra 
with fringe of long papillae on external border. Ely tral micro tubercles 
numerous, close-set, rather low, flattened, semiglobose, some bifid; 



218 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

macrotubercles confined mostly to a single row near external border, 
dark colored to pale yellow, nodular, with roughened tips or a fasicle 
of short spikes. Elytra often covered with a good deal of debris and 
foreign material. Color: In life and in alcohol: Mediodorsal 
surface colorless or banded with olive-brown between elytrophores and 
dorsal tubercles; ventral surface without color or olive-brown; setae 
yellow or light amber-colored; elytra yellow or tannish mottled with 
reddish brown. 

Remarks. — Eunoe nodosa has been confused with E. oerstedi. The 
two species have been separated by Sars (1860), Malmgren (1865), 
Verrill (1881), Murdoch and Benedict (1885, as Polynoe islandica and 
P. scahru), and Treadwell (1937). They have been considered to be 
synonymous and a highly variable species by Theel (1879), Wiren 
(1883), Fauvel (1923), Ditlevsen (1917), Augener (1928), and Wesen- 
berg-Lund (1950a, b). Based on a study of the material from Point 
Barrow as well as numerous other specimens in the U. S. National 
Museum from Greenland and off New England, the two species appear 
to be separable on the basis of a number of characters as indicated 
in the key. They agree in having the prostomium with cephalic 
peaks short and blunt or lacking, the position of the eyes, the dorsal 
cirri with long papillae, and the palps each with six longitudinal rows 
of papillae. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, stony-mud; off Point Barrow base, up to 12.1 miles 
from shore, 18.3-87 fathoms, on various combinations of mud, stones, 
pebbles, gravel, rocks (14 stations, 33 specimens). Bering Sea: 57° 
N., 163° 48' W., 38 fms., Alaska King Crab Expedition (Hartman 
(1948), as E. depressa); Albatross Sta. 3252, 57°22' N., 164 °24' W., 
29.5 fms., black mud, 1890, and Sta. 3512, 57°49' N., 169°27' W., 
38 fms., 1893. Southwestern Alaska: Belkofsky Bay, 15-25 fms., 
Dall, 1880. Franz Josef Land: Aberdore Chaimel, east Alger 
Island, 10 fms., Baldwin-Ziegler Expedition, 1901. East Coast 
North America: Off Labrador, 10-95 fms., Blue Dolphin Expeditions, 
1949, 1950; Albatross Sta. 2432, off Newfoundland, 43°04' N., 50°45' 
W., 64 fms., 1885; Bay of Fundy, Grand Manan, Nova Scotia, Maine, 
Massachusetts, 16-120 fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Baffin Bay, Davis Strait, Greenland, Jan 
Mayen, Spitsbergen, Norway, Franz Josef Land, Barents Sea, Novaya 
Zemlya, Kara Sea. Also Bering Sea; north Japan Sea; Iceland, Faroes, 
Shetlands to English Channel; Hudson Bay to Massachusetts. In 
10-690 fathoms. 



MARINE POLYCHAETE WORMS — PETTIBONE 219 

Eunoe oerstedi Malmgren, 1865 

Figure 26, d 

Lepidonote scabra Oersted, 1843, p. 164, pi. 1, figs. 2, 7, 10, 12-13, 17-18 (not 

Aphrodita scabra Fabricius, 1780). 
Polynoe scabra Sars, 1860, p. 58.— Hansen, 1882, p. 11.— Wir6n, 1883, p. 387, 

pi. 28, fig. 1 (part).— Murdoch, 1885, p. 152 (not A. scabra Fabricius, 1780). 
Eunoe oerstedi Malmgren, 1865, p. 61, pi. 8, fig. 3. — Moore, 1909b, p. 135. — 

Sumner, 1913, p. 618.— Treadwell, 1937, p. 27.— Hartman, 1944a, pp. 334, 

337.— Gorbunov, 1946, p. 38.— Pettibone, 1953, p. 46, pi. 23. 
Eunoe barbata Moore, 1910, p. 334, pi. 28, figs. 1-6.— Treadwell, 1925, p. 1.— 

Hartman, 1939, p. 53.— Berkeley and Berkeley, 1945, p. 321; 1948, p. 14, 

fig. 16.— Pettibone, 1949, p. 2. 
Eunoe nodosa Fauvel, 1923, p. 51, fig. 18, a-e (part). — Hartman and Reish, 1950, 

p. 7 (not Polynoe nodosa Sars, 1860). 
Harmothoe nodosa Ditlevsen, 1917, p. 6, pi. 2, fig. 1 (part; not pi. 3, fig. 10). — 

Augener, 1928, p. 684 (part) .— Wesenberg-Lund, 1950a, p. 6 (part); 1950b, 

p. 17 (part); 1951, p. 10 (part; not P. nodosa Sars, 1860). 

Description. — Length 18-73 mm., width including setae 6-29 mm. 
Segments 37-39 (39-42 in specimens from Washington). Prostomium 
with cephahc peaks poorly developed or lacking; anterior pair of eyes 
anterolateral, visible dorsally. Antennae and dorsal cirri with long 
papillae. Body elongate, oval in cross section. Segmental papillae 
short and bulbous. Tips of notosetae may be worn down and blunt, 
especially in larger specimens. Elytra with lateral fringe of papillae. 
Elytral surface exceedingly scabrous, with microtubercles one- to 
many-pronged; macro tubercles branched, extremely variable in size, 
number, arrangement and shape; some brownish, some translucent. 
Color: In alcohol : Dusky, dark or greenish black between elytrophores 
and dorsal tubercles, with some color on cirrophores of dorsal cirri 
and tips of parapodia; setae dark amber-colored. 

Remarks. — This species is closely related to and has been confused 
with E. nodosa as discussed above. Examination of the type of E. 
barbata Moore from central California revealed no essential differences 
from E. oerstedi. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, up to 8 miles from 
shore, 18.3 to 75.5 fms., on bottoms of mud, sand, shells, pebbles, 
gravel, rocks, stones (10 stations, 20 specimens). Canadian Arctic: 
Baffin Island, 66°43' N., 80°07' W., 1927; center, south, and southeast 
corner Foxe Basin, 25-37 fms., 1927; Kneeland Bay in Frobisher 
Bay, Baffin Island, 7-12 fms., 1942; all collected by Bartlett. West 
Greenland: Murchison Sound, 45-60 fms., Bartlett, 1938. East 
Greenland: Clavering Fiord, 1939; off Cape Hold with Hope, 
23-40 fms., Bartlett, 1939. South Greenland: Off Cape Farewell, 
70 fms., Bartlett, 1939. East Coast North America: Off Labrador, 

261112—54 2 



220 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

25-40 fms., Blue Dolphin Expeditions, 1949, 1950; Nova Scotia, 
Grand Manan, Maine, Massachusetts, low water to 110 fms., U. S. 
Fish Commission. Japan: Albatross Sta. 3656, Hakodate Bay, 11.5 
fms., 1896. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan 
and Canadian Arctic, Baffin Bay, Davis Strait, Greenland, Spits- 
bergen. Also from Norway to English Channel; Labrador to 
Massachusetts; Bering Sea to central California; and Japan. In 
low water to 516 fathoms. 

Genus Harmothoe Kinberg, 1857 

As used herein, Harmothoe includes Lagisca Malmgren, 1865; 
Evarne Malmgren, 1865; and Evarnella Chamberlin, 1919. 

Both species represented have the body short, attenuated posteriorly, 
segments about 40 (36-50), covered by elytra except for posterior 
few segments; elytra with numerous conical micro tubercles. Pro- 
stomium with cephalic peaks prominent (fig. 26, a). Antennae and 
dorsal cirri with short papillae. Notosetae stouter than neurosetae, 
with long distal spinous regions and short, pointed, bare tips. 

Key to the species of Harmothoe from Point Barrow 

1. Anterior pair of eyes antero ventral on prostomium (slightly posterior and 
lateral to cephalic peaks), not visible dorsally (fig. 26, a). Elytra with 
or without large, globular macrotubercles near posterior border. 

H. imbricata 
Anterior pair of eyes anterolateral on prostomium, visible dorsally. Elytra 
with macrotubercles globular, sausage-shaped or elongate, rodlike. 

H. extenuata 

Harmothoe imbricata (Linn^, 1767) 

Figure 26, a, e 

Aphrodita imbricata Linn6, 1767, p. 1804. 

Lepidonote cirrata Oersted, 1843, p. 166, pi. 1, figs. 1, 5-6, 11, 14-15. 

Polynoe cirrata Sars, 1860, p. 60. 

Harmothoe imbricata Malmgren, 1865, p. 66, pi. 9, fig. 8. — Webster and Benedict, 
1884, p. 701; 1887, p. 709.— Johnson, 1897, p. 181; 1901, p. 390.— Mcintosh, 
1900, p. 314, figs.— Moore, 1902, p. 270; 1903, p. 402; 1908, p. 334.— Dit- 
levsen, 1909, p. 7, pi. 2, fig. 5.— Sumner, 1913, p. 617.— Southern, 1914, p. 
52.— Chamberhn, 1920, p. 4.— EHason, 1920, p. 19.— Fauvel, 1923, p. 55, 
fig. 18, f-e; 1933, p. 10; 1936, p. 50.— Treadwell, 1925, p. 1; 1937, p. 26.— 
Augener, 1928, p. 677; 1939, p. 133.— Annenkova, 1934, p. 322; 1937, p. 151; 
1938, p. 136.— Okuda, 1938b, p. 83; 1939, p. 224.— Friedrich, 1939, p. 122.— 
Monro, 1939a, p. 345.— Hartman, 1944a, pp. 335, 337; 1948, p. 13.— Thorson, 
1946, p. 46, figs. 17-18.— Gorbunov, 1946, p. 38.— Zatsepin, 1948, p. 108, 
pi. 28, fig. 10.— Berkeley and Berkeley, 1948, p. 11, ^fig. 9.— Pettibone, 1949, 
p. 2; 1953, p. 32, pis. 13-16.— Hartman and Reish, 1950, p. 6.— Wesenberg- 
Lund, 1950a, p. 6; 1950b, p. 18; 1951, p. 12. 



MARINE POLYCHAETE WORMS — PETTIBONE 221 

Polynoe imbricata Marenzeller, 1879, p. 9, pi. 2, fig. 1. — Th6el, 1879, p. 9. — Hansen, 

1882, pp. 11, 13, 15, 23.— Wiren, 1883, p. 389. 
Harmothoe levis Treadwell, 1937, p. 26, figs. 1-5. 

Description. — Length 24-53 mm., width includmg setae 8-19 mm. 
(length up to 65 mm. — Theel, 1879). Segments 37-39. Prostomium 
with anterior pair eyes ventral, slightly posterior and lateral to 
cephalic peaks. Elytra with scattered conical microtubercles, with 
or without few to numerous, small, brownish to reddish, globular to 
elongate cylindrical macrotubercles in one, two, or several irregular 
rows near posterior border; may be a few additional globular papillae 
in middle of elj^'tra; with or without short elytral fringe of papillae. 
Neurosetae with long spinous regions, with long, bare, hooked tips, 
and usually with a subterminal tooth (may be broken off). Color: 
In life and in alcohol: Irregularly pigmented middorsally, on cirro- 
phores, dorsal tubercles, and parapodial lobes; usually without color 
ventrally; elytra show remarkable color variations of mottled tan and 
brown, blackish to grayish, or reddish; inner halves of elytra red or 
graj'^ish brown resulting in a middorsal, longitudinal, pigmented band ; 
uniformly tan; tan with bilateral darker spots near places of attach- 
ment (the color variet}^ of H. levis Treadwell) . 

Remarks. — Examination of the types of H. levis Treadwell from. 
Greenland waters revealed it to be but a color variation of H. 
imbricata. 

Parasites. — Of the 215 specimens, 13 had parasitic copepods, 
Herpyllobius arcticus Steenstrup and Liitken, attached on the pro- 
stomium (identified by Paul L. Illg). 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms., gravel, stones, mud; Point Barrow base, 
washed ashore; off Point Barrow base, up to 16 miles from shore, 
11.7-123.5 fms., on bottoms of stones, mud, masses of worm tubes, 
and various combinations of mud, pebbles, gravel, stones, large 
perforated rocks, with barnacles, bryozoans, hydroids, tunicates, 
shells, worm tubes (34 stations, 215 specimens). Canadian Artic: 
Southeast corner Foxe Basin, 66°46' N., 79°15' W., 34-37 fms., 1927; 
shoal in Kneeland Bay, Frobisher Bay, Baffin Island, 7-12 fms., 
1942; west shore Frobisher Bay, 1942; off Daniels Island, northwest 
side in Newell Sound, Frobisher Bay, 10-30 fms., 1942; all collected 
by Bartlett. West Greenland: Oelricks Bay, mud, 1937; Vaigat, 
Disko Island, mud, 1937; between Capes Alexander and Chalon, 
25-40 fms., rocks, 1937; Walrus grounds, Murchison Sound, 45-60 
fms., 1938; off Conical Rock, 76° N., 67°30' W., 20-40 fms., rocks, 
shell, 1938; all collected by Bartlett. Upernivik Harbor, 13 fms., 
U.S.S. Alert, 1884. East Greenland: Clavering Fiord, 1939; 
Angmogssalik, 10-15 fms., 1939; off Cape Hold with Hope, 23-40 
fms., 1939; all collected by Bartlett. Spitsbergen: Spitsbergen Sea, 



222 PROCEEDINGS OF THE NATIONAL MUSEUM vol.ios 

U.S.S. Alliance, 1881; South Gatt, northwest Spitsbergen, 79°40' N., 
7 fms., E. Wilkinson. East Coast North America: Off Labrador 
and Newfoundland, intertidal to 60 fms., Blue Dolphin Expeditions, 
1949, 1950, 1951; off Nova Scotia, Grand Manan, Maine, Massachu- 
setts, Rhode Island, Long Island Sound, intertidal to 110 fms., U. S- 
Fish Commission. Bering Sea: Bering Strait, 13 fms., Dall, 1880; 
62°54' N., 166°38' W., 22 fms., and 66°12' N., 168°54' W., Stoney, 
1884; St. George Island, Pribilofs, G. D. Hanna, 1914; Albatross 
Sta. 3252, 57°22' N., 164°24' W., 29.5 fms., 1890, and Sta. 3522, 
57°58' N., 170°09' W., 41 fms., 1893. Southwestern Alaska: Bay 
of Islands, Adak Island, 9-16 fms., 1873, and Coal Harbor, Unga 
Island, 1872, Dall. Southeastern Alaska: Wrangel, W. H. Jones. 
Sitka, "worm-eaten" wood, L. A. Beardslee. Japan: Albatross Sta. 
3656, Hakodate Bay, 1896. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Jan Mayen, Spits- 
bergen, Franz Josef Land, Barents Sea, Novaya Zemlya. Also 
Iceland and Norway to Mediterranean and Adriatic; Labrador to 
New Jersey; Bering Sea to southern California; and Japan. In low 
water to 2,030 fathoms. 

Harmothoe extenuata (Grube, 1840) 

Polyno'e extenuata Grube, 1840, p. 86 {fide Claparfede, 1868). — ClaparMe, 1868, 

p. 380, pi. 2, fig. 2. 
Polynoe rarispina Sars, 1860, p. 60.— Th^el, 1879, p. 8.— Hansen, 1882, p. 7.— 

Wir^n, 1883, p. 389. 
Lagisca rarispina Malmgren, 1865, p. 65, pi. 8, fig. 2. — Verrill, 1881, pp. 311, 

314.— Webster and Benedict, 1884, p. 700; 1887, p. 709.— Moore, 1902, p. 

269; 1908, p. 335.— Fauvel, 1911, p. 9; 1914, p. 65.— Chamberlin, 1920, p. 

5.— Berkeley and Berkeley, 1943, p. 129; 1948, p. 16, figs. 18, 19. 
Lagisca propinqua Malmgren, 1867, p. 9, pi. 1, fig. 3, a-e. 
Lagisca extenuata Marenzeller, 1876, p. 133, pi. 1, fig. 1; 1904, p. 318. — Mcintosh, 

1900, p. 307, figs.— Fauvel, 1923, p. 76, fig. 28, a-m; 1934a, p. 14.— Monro, 

1939a, p. 345.— Wesenberg-Lund, 1939a, p. 6, fig. 2; 1950a, p. 8; 1950b, p. 

27; 1951, p. 18. 
Lagisca impatiens Webster, 1879b, p. 102; 1886, p. 129, pi. 4, figs. 1-7. 
Polynoe semisculpta Hansen, 1882, p. 26, pi. 3, figs. 16-20 (?P. semisculptus 

Johnston, 1865). 
Lagisca fioccosa Mcintosh, 1900, p. 298, figs. — Southern, 1914, p. 51 (not Polynoe 

floccosa Savigny, 1820). 
Harmothoe extenuata Alaejos y Sanz, 1905, p. 55, pi. 9, fig. 8; pi. 10, figs. 1-12; 

pi. 11, fig. 1.— Ehlers, 1913, p. 446.— Pettibone, 1953, p. 31. 
Harmothoe rarispina Ditlevsen, 1909, p. 5, pi. 1, figs. 2-4; 1937, p. 11. — Augener, 

1928, p. 685.— Annenkova, 1937, p. 153; 1938, p. 136.— Friedrich, 1939, p. 

122.— Gorbunov, 1946, p. 38.— Zatsepin, 1948, p. 108, pi. 28, fig. 15. 
Harmothoe triannulata Moore, 1910, p. 346, pi. 29, figs. 18-22. — Hartman, 1938a, 

p. 118.— Berkeley and Berkeley, 1948, p. 12, fig. 10. 
Lagisca extenuata var. spinulosa Fauvel, 1914, p. 64, pi. 4, figs. 27-29. 
Harmothoe propinqua Ditlevsen, 1917, p. 14, pi. 3, figs. 1, 3. 
Harmothoe rarispina forma propinqua Annenkova, 1937, p. 153; 1938, p. 137. 



MARINE POLYCHAETE WORMS — PETTIBONE 223 

Evarnella triannulata Berkeley and Berkeley, 1942, p. 188; 1943, p. 130. — Hartman, 
1948, p. 13 (part; includes H. muliisetosa Moore and H. extenuata). — Petti- 
bone, 1949, p. 1. 

Description. — Length 13-68 mm., width including setae 4-20 mm. 
(length up to 74 mm. — Ditlevsen, 1937). Segments 37-47. Prosto- 
mium with eyes large, anterior pair anterolateral, slightly anterior to 
widest part of prostomium, visible dorsally. Neurosetae with en- 
larged, long spinous regions, with tips slightly hooked, with small, 
secondary tooth present or absent; at least some of the setae show a 
remnant of a secondary tooth. The upper and lower neurosetae tend 
to be unidentate (not always) while the middle ones are bidentate, 
with a secondary tooth or remnant of it (in specimens from Woods 
Hole region only a few neurosetae have a secondary tooth or remnant 
of one). Elytra with numerous micro tubercles, more or less uniform 
in size, conical, mth tips blunt, pointed, or bifid; usually with few to 
fairly numerous macrotubercles distinctly set off from elytral surface, 
usually narrower at the base, brown, smooth, globular, sausage- 
shaped, or elongate fusiform, variable in number, 0-9 near posterior 
border, 0-13 scattered near center of elytron. Elytra with short 
fringe of papillae on external border. Color: In alcohol: Without 
color anteriorly; on posterior half, with brownish transverse bands and 
two brown spots basallj'' on cirrophores (in specimens from Woods 
Hole region, found intertidally, the body is darldy pigmented grayish- 
green middorsally) ; antennae and dorsal cirri ringed with brown; 
elytra mottled with brown, sometimes with a darker spot posterior 
and medial to the place of attachment; the micro tubercles may be 
covered with a brown extraneous material, giving a streaked appearance. 

Remarks. — H. extenuata is an extremely variable species as indicated 
by the number of names that have been applied to it. It reaches its 
greatest size in arctic waters; it is small when found intertidally, as 
in the Puget Sound region (up to 23 mm. — H. triannulata of Moore). 
When found intertidally off New England, it also is small (about 26 
mm.), and the secondary tooth of the neurosetae is suppressed, usually 
with only a remnant showing on a few neurosetae, the rest being entire 
(L. impatiens of Webster). In the collection from Point Barrow, 
where it was the most abundant polychaete, it is represented by 
specimens of all sizes from very small to large (up to 68 mm.). 

Parasites. — Of the 375 specimens, 3 had the parasitic copepod 
Herpyllohius arcticus Steenstrup and Lutken attached to the pros- 
tomium (identified by lUg) . 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, up to 15 miles from 
shore, 13.3-123.5 fms., on bottoms of mud, stones, masses of worm 
tubes, and various combinations of mud, pebbles, rocks, gravel, 
stones, large perforated rocks, with shells, bryozoans, hydroids, 



224 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

worm tubes, from starfish and crab, Hyas coarctatus (37 stations, 375 
specimens). Canadian Arctic: Foxe Basin, 66°30' N., 80° W., 
1927; center Foxe Basin, 25-31 fms., 1927; 7 miles east Cape Dor- 
chester, Foxe Channel, 25 fms., 1927; Baffin Island, 66°43' N., 80°07' 
W., 1927; Hurd Channel between Bushman Island and Melville 
Peninsula, 11 fms., rocky, 1933; south end Cape Martineau, Melville 
Peninsula, 7-15 fms., 1933; 3 miles south Salisbury Island, Hudson 
Strait, 27 fms., 1933; south end Cobourg Island, Baffin Bay, 75°40' 
N., 78°58' W., 11.3-20 fms., 1935; east end Cobourg Island, 75°40' 
N., 78°50' W., 23-36 fms., 1935; southeast corner Foxe Basm, 
66°46' N., 79°15' W., 34-37 fms., 1937; Kneeland Bay, Frobisher 
Bay, Baffin Island, 7-12 fms., 1942; all collected by Bartlett. West 
Greenland: Vaigat, Disko Island, mud, 1937; between Capes 
Alexander and Chalon, 25-40 fms., rocky, 1937; Walrus grounds, 
Murchison Sound, 45 fms., 1938; all collected by Bartlett. North 
Greenland: North Omenolu near North Star Bay, 17 fms., and Cape 
Alexander, entrance to Smith Sound, rocky, 1932, Bartlett. East 
Greenland: Off Cape Hold with Hope, 23-40 fms., Bartlett, 1939. 
South Greenland: Cape Farewell, 70 fms., Bartlett, 1939. Franz 
Josef Land: Aberdore Channel east Alger Island, 10 fms., Baldwin- 
Ziegler Polar Expedition, 1901. Bering Sea: Alaska King Crab 
Investigation, 1941, Sta. D8-41, 58°34' N., 165°17' W., 42 fms.; 
Sta. D7-41, 57° N., 163°48' W., 38 fms.; Sta. Dll-41, 12 miles east 
Walrus Island, Pribilofs, 31-33 fms.; St. George Island, PribUofs, 
village beach, 1914, and St. Paul Island, Pribilofs, 1915, Hanna; 
62°54' N., 166°38' W., 22 fms., and 62°15' N., 167°48' W., 20 fms., 
Stoney, 1884. Southwestern Alaska: Alaska King Crab Investiga- 
tion, 1940, 1941, Sta.: 89-40, Unga Strait, 37-47 fms.; 93-40, Spitz 
Island, 55-68 fms.; Sand Point; 128-40, Shelikof Strait off Hallo Bay, 
35-48 fms.; 60-40, Lenard Harbor, 20-25 fms.; L18-41, Kupreanof 
Strait, south side, 2 miles northwest Bare Island, 13-15 fms.; 72-40, 
Cold Bay, 15-50 fms.; 52-40, Canoe Bay, 35-40 fms., on Paralithodes 
camtschatica; 100-40, Alitak Bay, 30 fms.; 59-40, between inner 
Iliasik and Goloi Island, 20-30 fms. Southeastern Alaska: 
Lituya Bay, 6-9 fms., Dall, 1874. East Coast North America: 
Off Labrador, 6-100 fms.. Blue Dolphin Expeditions, 1949, 1950, 
1951; Bay of Fundy, Gulf of Maine, Massachusetts, Long Island 
Sound, 4-134 fms., U. S. Fish Commission; Woods Hole region, 
intertidal and dredged, Pettibone, 1950, 1951; Fish Hawk Sta. 8826, 
8898, Chesapeake Bay, 1920. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Ai-ctic, Baffin Bay, Davis Strait, Greenland, Spits- 
bergen, Franz Josef Land, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes, Norway to Mediterranean and Adriatic; Hudson Bay to 
Chesapeake Bay; Bering Sea to southern California; north Japan Sea; 
South Africa. In low water to 1,000 fathoms. 



MARINE POLYCHAETE WORMS — PETTIBONE 225 

G^nus Enipo Malmgren, 1865, sensu Levinsen, 1883 
Enipo gracilis Verrill, 1874 

Enipo gracilis VerriU, 1874a, pp. 407, 411, pi. 6, fig. 4; 1874b, pp. 347, 361, 378, 

pi. 5, fig. 3.— Pettibone, 1953, p. 22, pi. 7. 
Enipo cirrata Treadwell, 1925, p. 1, figs. 1-4. — Hartman, 1938a, p. 120, &g. 39, 

d, e.— Berkeley and Berkeley, 1942, p. 188; 1945, p. 322; 1948, p. 16, fig. 20. 
Polynoe gracilis Hartman, 1942a, p. 26, figs. 23-26, 

Description. — Length 25-76 mm., width including setae 3-4 mm. 
Segments variable in number (45, with regenerating posterior end, 
to 74). Body elongate, slender, with sides nearly parallel. Prosto- 
mium without cephalic peaks. In some specimens from New England 
there are slight prostomial peaks (prostomial peaks present — Hart- 
man, 1942). Antennae and dorsal cirri with short, clavate papillae. 
Dorsal cirri short, barely reaching tips of setae in anterior part, longer 
in posterior region without elytra. Elytra oval, translucent, small, 
usually leaving middorsum uncovered, smooth, without papillae or 
tubercles. Notosetae form a short, bushy bundle, rather numerous 
(more than 30 in specimens from Point Barrow) to few (2-20 or 
rarely — Treadwell, 1925), slender to moderately stout, tapering 
gradually to short, bare, blunt tips, with spinous rows distally. 
Neurosetae stouter than notosetae, upper ones with longer spinous 
regions (17-23 or so rows), middle ones with shorter spinous regions 
(10 or so rows), with long, bare, slightly hooked tips (neurosetae may 
be all similar, with 5-10 transverse spinous rows) . Color : In alcohol : 
Without color or brownish middorsally; elytra pigmented smoky 
brown on the medial halves. 

Remarks. — -Known to be commensal with the maldanid Nicomache 
lumhricalis (Fabricius) in Alaska (as E. cirrata — Berkeley and Berke- 
ley, 1942) and off Halifax, Nova Scotia, and Cape Cod (specimens in 
the U. S. National Museum, identified by Verrill). 

New Records. — Arctic Alaska: Off Point Barrow, up to 12.1 miles 
from shore, 21-123.5 fms., from breaking rock, masses of worm tubes, 
with stones, large perforated rocks (3 stations, 3 specimens). East 
Coast North America: Off Nova Scotia, Maine, Massachusetts, 2- 
198 fms., U. S. Fish Commission. 

Distribution. — Alaskan Arctic; southwestern Alaska to Washington; 
Nova Scotia to Massachusetts. In 2-123.5 fathoms. 

Genus Arcteobia Annenkova, 1934 
Arcteobia anticostiensis (Mcintosh, 1874) 

Eupolynoe anticostiensis Mcintosh, 1874, p. 265, pi. 10, figs. 1-4. — Wir6n, 1883, 

p. 390, pi. 29, fig. 2.— Marenzeller, 1890, pi. 1. 
Harmothoe anticostiensis Augener, 1928, p. 691, pi. 11, fig. 13. 



226 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Ardeobia anticostiensis Annenkova, 1934, p. 322; 1937, p. 149, pi. 3, figs. 26, 27; 

1938, p. 133. 
Eucranta anticostiensis Hartman, 1944a, p. 337. 

Description. — Length 11-26 mm., width mcluding setae 4-8 mm. 
Segments 35 or 36. Prostomium with distinct cephaHc peaks; anterior 
pair of eyes antero ventral, not visible dorsally. Antennae and dorsal 
cirri with short, clavate papillae. Elytral pairs get larger posteriorly, 
the last pair usually extending beyond end of body. Elytra without 
fringe of papillae, smooth except for scattered microtubercles on 
anterior curved part. Upper notosetae shorter, stouter, with short, 
blunt to sharp-pointed tips; most of notosetae more slender, with 
capillary tips. Few upper neurosetae longer, ending in sharp, slender 
tips;mostofnem'Osetae with bifid tips. Color: In life and iu alcohol : 
Irregularly banded middorsally, gi-eenish to greenish black, with a 
transverse stripe and often a pair of small spots; elytra with reddish 
brown pigmented area, usually C-shaped on inner and posterior parts 
and often a darker spot over place of attachment; may form a com- 
plete, circular, colored area. 

Remarks.- — -Probably commensal in habit; off Labrador, found com- 
mensal in the sinuous tubes of the terebeUid Pista Jlexuosa (Grube), 
one worm per tube. 

New records. — Arctic Alaska: Point Barrow base, washed ashore; 
off Pomt Barrow base, up to 12.1 miles from shore, 24.7-123.5 fms., 
on bottoms of mud, masses of worm tubes, and various combinations 
of mud, gravel, stones, rocks, and large perforated rocks (7 stations, 
14 specimens). East Coast North America: Off Labrador, 40-95 
fms.. Blue Dolphin Expedition, 1949; off Nova Scotia, Maine, Mas- 
sachusetts, 10-86 fms., U. S. Fish Commission. 

Distribution. — Alaskan Arctic; Labrador to Massachusetts; Bering 
Sea; north Japan Sea. In low water to 123.5 fathoms. 

Genus Gattyana IVIcIntosh, 1897 

Key to the species of Gattyana from Point Barrow 

1. Notosetae with tips blunt to capillary. Anterior pair of eyes anteroventral, 
not visible dorsally; cephalic peaks distinct. Elytra with microtubercles 
only G. cirrosa 

AH notosetae with capillary tips. Anterior pair of eyes anterolateral; cephalic 
peaks short and blunt. Elytra with conical and bifid microtubercles, with a 
few larger, knobUke macrotubercles G. ciliata 

Gattyana cirrosa (Pallas, 1766) 

Figure 26, b 

Aphrodita cirrhosa Pallas, 1766, p. 95, pi. 8, figs. 3-6. 

Nychia cirrosa Malmgren, 1865, p. 58, pi. 8, fig. 1.— Th6el, 1879, p. 7.— VerriU, 

1881, pp. 306, 311.— Wir6n, 1883, p. 387.— Webster and Benedict, 1884, p. 

700; 1887, p. 708.— ? Andrews, 1891, p. 279. 



MARINE POLYCHAETE WORMS — PETTIBONE 227 

Gattyana cirrosa Mcintosh, 1900, p. 285, figs.— Moore, 1902, p. 259; 1908, p. 
337.— Southern, 1914, p. 51.— EHason, 1920, p. 21.— Fauvel, 1923, p. 49, 
fig. 17, a-g.— Augener, 1928, p. 692; 1939, p. 136.— Gustafson, 1936, p. 5.— 
TreadweU, 1937, p. 25.— Annenkova, 1937, p. 148; 1938, pp. 83, 96, 133, 
222.— Monro, 1939a, p. 345.— Berkeley and Berkeley, 1943, p. 129; 1948, 
p. 13, fig. 13.— Thorson, 1946, p. 45.— Gorbunov, 1946, p. 38.— Hartman, 
1948, p. 14.— Zatsepin, 1948, p. 106, pi. 28, fig. 5.— Pettibone, 1949, p. 2; 
1953, p. 41, pi. 20.— Wesenberg-Lund, 1950a, p. 5, pi. 1, figs. 1, 2; 1950b, 
p. 12; 1951, p. 9. 

Gattyana cirrhosa Chamberlin, 1920, p. 8. 

Description. — Length 11-35 mm., width including setae 4-12 mm- 
Segments 35-38. Elytra with long papillae on posterior and external 
borders as well as scattered over the exposed part of elytral surface, 
usually covered with debris, giving a straggly appearance. Elytral 
tubercles amber-colored, exceedingly variable in shape, conical or 
cylindrical, simple, bifid or quadrifid, showing various degrees of 
being worn down; tubercles on the anterior few pairs of elytra tend 
to be less worn down, showing their more characteristic shapes. 
Color: In alcohol: Elytra tan mottled with brown or uniformly tan; 
often with blackish foreign material on elytral tubercles and papillae. 
Specimens washed ashore were essentially free from debris, the elytra 
whitish, iridescent, with amber-colored tubercles and inconspicuous, 
delicate papillae. 

Parasites. — Three of the 126 specimens had the parasitic copepod 
Herpyllobius arcticus Steenstrup and Liitken attached to the prostomia 
(identified by Illg). 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; Point Barrow base, washed ashore; off Point 
Barrow base, up to 15 miles from shore, 18.3-123.5 fms., on bottoms 
of mud, stones, masses of worm tubes, and various combinations of 
mud, pebbles, small and large rocks, stones, gravel, and worm tubes; 
from breaking rock (30 stations, 126 specimens). Cape Smyth, 2.5-3 
fms.. Point Barrow Expedition, 1883. Canadian Arctic: South end 
Cape Martineau, Melville Peninsula, 7-15 fms., 1933; Foxe Basin, 
1933; east end Cobourg Island, Baffin Bay, 75°40' N., 78°50' W., 
23-40 fms., 1935; Kneeland Bay, Frobisher Bay, Baffin Island, 14 
fms., 1942; all collected by Bartlett. West Greenland: Vaigat, 
Disko Island, 1937; between Capes Alexander and Chalon, 25-40 fms., 
rocky, 1937; Walrus grounds, Murchison Sound, 60 fms., 1938; 
northwest Conical Rock, 1938; all collected by Bartlett. Upernivik 
Harbor, 13 fms., and Godhavn, U. S. S. Alert, 1884. East Green- 
land: Off Cape Hold with Hope, 4-6 fms., Bartlett, 1939. North 
Greenland: North Omenolu near North Star Bay, 17 fms., Bartlett, 
1932. SpnsBERGEN: Spitsbergen Sea, U. S. S. ylZ^iance, 1881. Bering 
Sea: Bering Strait, 13 fms., Dall, 1880. Albatross Sta. 3326, 53°40' 



228 PROCEEDINGS OF THE NATIONAL MUSEUM tol. 103 

N., 167°41' W., 1890. Southwestern Alaska: Iluiliiik Harbor, 
Unalaska, Dall, 1871. Southeastern Alaska: Sitka, Dall. East 
Coast North America: Off Labrador, 13-125 fms., Blue Dolphin 
Expeditions, 1949, 1950, 1951. 

Distribution. — -Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Spitsbergen, Novaya 
Zemlya. Also Iceland, Faroes, Norway to France; Hudson Bay to 
Massachusetts; Bering Sea to Washington; north Japan Sea. In 
low water to 630 fathoms. 

Gattyana ciliata Moore, 1902 

Gattyana ciliata Moore, 1902, p. 263, pi. 13, figs. 14-19; pi. 14, fig. 20; 1905a, 
p. 525; 1908, p. 337.— Annenkova, 1937, p. 148, pi. 2, figs. 9, 11; pi. 4, fig. 33; 
1938, pp. 83, 100, 132, 224.— Berkeley and Berkeley, 1948, p. 12, fig. 11.— 
Pettibone, 1953, p. 40, pi. 19. 

Description.- — Length 63-65 mm., width including setae 25-29 mm, 
(length up to 80 mm.^ — ^Annenkova, 1937). Segments 36 or 37. 
Cephalic peaks lacking or short and blunt. Elytral tubercles of 
several kinds: Microtubercles numerous, conical, pointed, hooked, 
some bifid; usually intermediate-sized tubercles elongated, conical, 
some bifid, in several rows arranged diagonally from center of elytron 
lateralljT^; conical macro tubercles Vv^ith blunt, roughened tips, usually 
near posterior border of elytra, variable in number, size, and shape. 
Color: In alcohol: Elytra tannish mottled with brown. 

New records. — Arctic Alaska: Oft' Point Barrow base, up to 12.1 
miles from shore, 20-87 fms., on bottoms of rocks, stones, smaU 
amount of gravel (4 stations, 4 specimens). Bering Sea: Albatross 
Sta. 3606, 55°27' N., 167°47' W., 87 fms., green mud, fine sand, 1895. 

Distribution. — Alaskan Arctic (originally recorded from Greenland 
waters, corrected to Icy Cape, Alaska — Moore, 1902, 1905); Bering 
Sea to Washington; north Japan Sea. In 4.4-303 fathoms. 

Family Sigalionidae 

Prostomium subglobular, with two pairs of eyes, dorsal antennae 
1-3, a pair of subulate, ventral palps (fig. 26,/). First or tentacular 
segment with or without numerous setae (lacldng in Pholoe), with 
two pairs of tentacular cirri. Parapodia biramous, with notosetae 
simple, neurosetae simple or composite or both; with paired dorsal 
elytra on certain segments ; without dorsal cirri ; with paired subulate 
ventral cirri. Pygidium with pair of anal cirri. With or without 
cirriform branchiae and ciliated cushions or ctenidia on parapodia 
(both lacking in Pholoe) . Muscular proboscis eversible, with circle of 
soft marginal papillae and two pairs of interlocking, chitinous jaws. 

Represented by a single species from Point Barrow. 



MARINE POLYCHAETE WORMS — PETTIBONE 




Figure 26. — Polynoidae: a, Harmothoe imbricata, dorsal view anterior end, first pair elytra 
removed; h, Gattyana cirrosa, part of elytron enlarged, showing elytral tubercles and 
papillae; c, Euno'e nodosa, dorsal view part of two segments showing extra rounded lobes 
on inner sides of elytrophore and dorsal tubercle; d, Eunoe oerstedi, biramous parapodium; 
Cy Harmothoe imbricata, bidentate neuroseta. Sigalionidae; /, Pholoe minuta, dorsal 
view anterior end, first pair elytra removed. (For explanation of symbols, see p. 210.) 



230 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m 

Genus Pholoe Johnston, 1839 
Pholoe minuta (Fabric! us, 1780) 

Figure 26, / 

Aphrodita minuta Fabricius, 1780, p. 314. 

Aphrodila longa Fabricius, 1780, p. 313. 

Pholoe minuta Oersted, 1843, p. 169, pi. 1, figs. 3, 4, 8, 9, 16. — Malmgren, 1865, 
p. 89, pi. 11, fig. 13.— Th6el, 1879, p. 24.— Webster and Benedict, 1884, 
p. 701; 1887, p. 709.— Mcintosh, 1900, p. 437, figs.— Moore, 1902, p. 274; 
1908, p. 338. — Ehlers, 1913, p. 450.— Southern, 1914, p. 57. — Chamberlin, 
1920, p. 5.— Eliason, 1920, p. 22.— Fauvel, 1923, p. 120, fig. 44, a-h.— 
Augener, 1928, p. 673.— Annenkova, 1934, p. 322; 1937, p. 155; 1938, p. 138.— 
Berkeley and Berkeley, 1942, p. 189.— Hartman, 1944a, pp. 335, 337; 1948, 
p. 15.— Thorson, 1946, p. 49, fig. 19; Gorbunov, 1946, p. 38.— Zatsepin, 
1948, p. 109, pi. 28, fig. 17. — Hartman and Reish, 1950, p. 8.— Wesenberg- 
Lund, 1950a, p. 9; 1950b, p. 31; 1951, p. 22.— Pettibone, 1953, p. 77, pi. 39. 

Pholoe tuberculata Southern, 1914, p. 57, pi. 6, fig. 14, A-L. — Berkeley and 
Berkeley, 1945, p. 323; 1948, p. 22, fig. 26. 

Description. — Length 5-25 mm., width including setae 1-4 mm. 
Segments 36-84. Body small, elongate, nearly linear, flattened 
dorsoventrally; fragments easily. Lateral ventral surface and ventral 
sides of parapodial lobes thickly set with small, globular to elongate 
papillae, usually covered with debris. Prostomium small, oval; 
anterior and posterior pairs of eyes closely approximated. Single 
median antenna short, subulate (lateral antennae lacking). Ten- 
tacular segment achaetous; tentacular cirri similar to median antenna. 
A digitiform facial tubercle on a rounded lobe dorsal to the mouth 
and ventral to the prostomium. (On this character, Southern (1914) 
differentiated P. tuberculata; the lobe, on which the facial tubercle is 
located, appears to be retractile; surrounded by the bases of the palps, 
tentacular cirri and median antenna, the facial tubercle is easily 
overlooked unless the region is quite relaxed.) Elytra numerous, on 
segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 24, continuing on all 
segments to end of body; elytra nearly cover the dorsum except for a 
narrow middorsal part, usually pressed close to body; elytral surface 
smooth, with few microtubercles, somewhat areolate as seen under 
magnification, posterior border with few scattered papillae, somewhat 
moniliform. Notosetae simple, slender, finely spinous, tapering to 
capillary tips, upper ones shorter, with pronounced angles, the rest 
longer, with more gradual curves. Neurosetae stouter than notosetae, 
compound, with short, falcate terminal pieces. Color: In alcohol: 
Without color or irregularly pigmented greenish gray; elytra trans- 
lucent, colorless, or greenish gray with a light spot over the place of 
attachment, or brownish; often with much debris. 



MARINE POLYCHAETE WORMS — PETTIBONE 231 

Nev) records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, up to 12.1 miles from 
shore, 16.7-123.5 fms., on bottoms of stones, masses of worm tubes, 
and various combinations of mud, gravel, stones, rocks, large per- 
forated rocks, with bryozoans, hydroids; from screen trap through 
hole in ice, mud (16 stations, 30 specimens). West Greenland: 
Vaigat, Disko Island, mud, 1937, and Walrus grounds, Murchison 
Sound, 45 fms., 1938, Bartlett. East Coast North America: Off 
Labrador, 6 fms.. Blue Dolphin Expedition, 1949; off Nova Scotia, 
St. Georges Bank, Maine, Massachusetts, Rhode Island, intertidal to 
110 fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Jan Mayen, Norwegian 
Sea, Spitsbergen, Franz Josef Land, Novaya Zemlya, Kara Sea. 
Also Iceland, Faroes, Shetlands, Norway to northern France ; Labrador 
to Rhode Island; Bering Sea to southern Oregon; north Japan Sea; 
off South Africa. In low water to 1,254 fathoms. 

Family Phyllodocidae 

Prostomium subcorneal, suboval or cordiform, with two eyes, four 
or five antennae, without palps (fig. 27, a-<i). Anterior segments 
1-3 modified, with two to four pairs tentacular cirri. Parapodia 
uniramous (exceptionally biramous); setae compound (some may be 
simple). Dorsal and ventral cirri foliaceous or more or less globular 
(fig. 27, e-4). Two anal cirri. Proboscis eversible, with papillae, 
unarmed. Active; mucus secreted in quantities. 

Represented by four genera and seven species. All genera repre- 
sented have uniramous parapodia, with compound setae. 

Key to the genera of PhyUodocidae from Point Barrow 

1. Tentacular cirri 4 pairs; first tentacular segment rudimentary, not visible 
dorsally, with 1 pair tentacular cirri lateral to prostomium; second segment 
distinct, with 2 pairs tentacular cirri; third segment distinct, with 1 pair 
tentacular cirri, 1 pair normal ventral cirri (fig. 27, c, d) 2 

Tentacular cirri 3 pairs; first tentacular segment distinct dorsally, with 1 pair 
tentacular cirri; second segment distinct, with 2 pairs tentacular cirri (fig. 
27, b). Prostomium oval, without occipital tubercle Mystides (p. 232) 

Tentacular cirri 2 pairs on first achaetous segment (fig. 27, a) . Without dorsal 
cirri on second segment which has a setigerous lobe (may be very small or 
lacking) and a foliaceous ventral cirrus. Prostomium somewhat triangular, 
widest basally, with anterior part rounded, with 2 pairs short, subulate, 
frontal antennae, usually with 2 deep-set eyes (may not be visible when 
preserved), usually with a median occipital depression containing a small 
occipital tubercle (not always conspicuous), with a pair of lateral nuchal 
grooves Eteone (p. 232) 



232 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

2. Antennae 4 (2 pairs frontal antennae). Prostomium cordiform, with an 
occipital tubercle in the posterior notch (fig. 27, d) . Proboscis with papillae 
proximally arranged in longitudinal rows Phyllodoce (p. 236) 

Antennae 5 (2 pairs frontal antennae and unpaired median antenna) . Prosto- 
mium oval or bluntly conical, without occipital tubercle (fig. 27, c). Pro- 
boscis with surface smooth, wrinkled, or with few scattered papillae 

Eumida (p. 238) 

Genus My slides Theel, 1879 
Mystides borealis Theel, 1879 

Figure 27, b 

Mystides borealis Thdel, 1879, p. 35, pi. 2, figs. 29-32.— Southern, 1914, p. 72, 
pi. 8, fig. 19, A-D.— Bergstrom, 1914, p. 176, fig. 64.— Fauvel, 1923, p. 181, 
fig. 65, a-d.— Augener, 1928, p. 711.— Zatsepin, 1948, p. 113, pi. 29, fig. 15.— 
Wesenberg-Lund, 1950b, p. 37. 

Mystides viridis Webster and Benedict, 1887, p. 712, pi. 1, figs. 10, 11, 13; pi. 2, 
fig. 12. (Type in USNM.) 

Mystides notialis Ehlers, 1913, p. 457, pi. 29, figs. 1-4. 

Description. — Length 5-9 mm., width 0.5 mm. Segments 33-64 
(up to 16 mm. long and 73 segments — Ehlers, 1913). Body small, 
linear, tapering anteriorly and posteriorly, flattened dorsoventrally. 
Prostomium ova], wider than long, with two deeply-set eyes in middle 
of prostomium, with four long, filiform antennae about one-third 
way back from anterior tip. Tentacular cirri 3 pairs, enlarged basally, 
with long, filiform tips. Dorsal, ventral, and anal cirri oval, thick, 
flattened. Color: In alcohol: Without color or irregularly pigmented 
with brown; cirri deep reddish brown. 

Remarks. — The description of M. notialis from the Antarctic by 
Ehlers (1913) follows closely that of M. borealis and is herein considered 
to be synonymous. 

New records. — ^Arctic Alaska: Off Point Barrow, 7.5 miles from 
shore, 36 fms., on bottom of stones, perforated rocks (1 station, 2 
specimens). West Coast North America: Off Flat Point, Lopez 
Island, Washington Sound, Pettibone (1 specimen). 

Distribution. — Scattered records in the Arctic: Arctic Alaska, West 
Greenland, Spitsbergen, Novaya Zemlya. Also Ireland, Madeira, 
Mediterranean; east coast North America (Maine); west coast North 
America (Washington Sound); Antarctic (Kerguelen). In 4-214 
fathoms. 

Genus Eteone Savigny, 1817 

The four species represented have the body elongate, somewhat 
flattened dorsoventrall}^, tapering anteriorly and posteriorly, with 
segments numerous (100-300). Proboscis with a circle of soft papillae 
around opening. 



MARINE POLYCHAETE WORMS — PETTIBONE 233 

Key to the species of Eteone from Point Barrow 

1. Proboscis with 2 lateral, longitudinal rows of numerous soft papillae. Dorsal 

pair tentacular cirri slightly longer or up to 2-3 times longer than ventral 
pair. Dorsal cirri wider than long, nearly symmetrical, subcircular or 
slightly lanceolate (fig. 27, e). Color in alcohol: Darkly pigmented with 
bluish-violet iridescence, with longitudinal, lighter colored bands on each 
side of median dorsal line (or 3 dorsal longitudinal dark bands — 1 median 
and 2 lateral). Anal cirri subcylindrical, 3-4 times longer than wide. 

Second segment with setigerous lobe and setae well developed E. barbata 

Surface of proboscis smooth, rugose or irregularly papillate. Two pairs ten- 
tacular cirri subequal or ventral pair somewhat longer than dorsal pair — 2 

2. Dorsal cirri small, not much larger than parapodial lobes, longer than or as 

long as wide, almost symmetrical, thick, flattened, bluntly conical (fig. 27, 
h). Anal cirri short, thick, almost spherical. Second segment with seti- 
gerous lobe and setae well developed E. longa 

Dorsal cirri much larger than parapodial lobes, v/ider than long 3 

3. Dorsal cirri slightly asymmetrical, oval (fig. 27, g). Color in alcohol: Dorsum 

brownish with bluish-violet irridescence. Anal cirri short, thick, almost 
spherical. Second segment with setigerous lobe and setae well developed. 

E. flava 
Dorsal cirri much wider than long (about 2.5 times), asymmetrical, obliquely 
oval, with lower part larger than upper part (fig. 27, /). Color in alcohol: 
Middle third of dorsum light brownish, lateral third deep reddish brown. 
Anal cirri elongate, cylindrical, 3-4 times longer than wide. Second segment 
with setigerous lobe and setae poorly developed or lacking. 

E. spetsbergensis 



Subgenus Mysta Malmgren, 1865 
Eteone {Mysta) barbata (IVIalmgren, 1865) 

Figure 27, e 

Mysta barbata Malmgren, 1865, p. 101, pi. 15, fig. 34; 1867, p. 26, pi. 3, fig. 20.— 
Bergstrom, 1914, p. 207, fig. 79.— Southern, 1914, p. 75.— Eliason, 1920, 
p. 28.— Augener, 1928, p. 711; 1939, p. 136.— Zatsepin, 1948, p. 113, pi. 29, 
fig. 14. — Wesenberg-Lund, 1951, p. 31. 

Eteone striata Ditlevsen, 1917, p. 66, pi. 5, figs. 11, 17, 19. 

Eteone (Mysta) barbata Fauvel, 1923, p. 176. — Annenkova, 1937, p. 159; 1938, 
p. 146.— Berkeley and Berkeley, 1942, p. 190.— Thorson, 1946, p. 62, fig. 
27, A-B. 

Description. — Length 42-180 mm.; width without setae 2.5-7 mm. 
See key. Color: In life: Three iridescent, brownish-purple dorsal 
longitudinal bands, median and lateral. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, stony-mud; off Point Barrow base, 6.1-27 fms., up 
to 3.2 miles from shore, on bottoms of mud, gravel, stones, rocks; 
also caught in trap on bottom (3 stations, 4 specimens). 



234 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Distribution. — Scattered records in the Arctic : Siberian and Alaskan 
Arctic, Spitsbergen, Novaya Zemlya, Kara Sea. Also Iceland, Nor- 
way to Ireland, North Sea, Baltic; Bering Sea; north Japan Sea. 
In 5.5-60 fathoms. 



Eteone longa (Fabricius, 1780) 

FiGUKE 27, h 

Nereis longa Fabricius, 1780, p. 300. 

Eteone longa Oersted, 1843, p. 185, pi. 2, figs. 20, 28. — Bergstrom, 1914, p. 192, 
fig. 72.— Chamberlin, 1920, p. 11.— Eliason, 1920, p. 26.— Fauvel, 1923, 
p. 172, fig. 62, a-d; 1933, p. 16.— Augener, 1928, p. 710; 1939, p. 136.— 
Annenkova, 1934, p. 322; 1937, p. 158; 1938, p. 145.— Thorson, 1946, p. 
59, fig. 26, A-E.— Gorbunov, 1946, p. 38.— Zatsepin, 1948, p. 113, pi. 29, 
fig. 13. — Berkeley and Berkeley, 1948, p. 41, figs. 57, 58. — Wesenberg-Lund, 
1950a, p. 11, pi. 2, figs. 6, 7; 1950b, p. 38; 1951, p. 29. 

Eteone arctica Malmgren, 1867, p. 27, pi. 2, fig. 12. 

Eteone cinerea Webster and Benedict, 1884, p. 705, pi. 1, figs. 1-5. 

Eteone tuberculata Treadwell, 1922, p. 174, figs. 7-10. 

Eteone californica Hartman, 1936a, p. 131, figs. 43-46; 1948, p. 20, fig. 4, a-d. — 
Rioja, 1941, p. 687. 

Description. — Length 10-65 mm., width without setae 0.5-3 mm. 
(length up to 160 mm. — Berkeley and Berkeley, 1948). See key. 
Color: In life: White. 

Remarks. — E. tuberculata Treadwell (1922) from Friday Harbor, 
Washington, E. calijornica Hartman (1936) from central California 
(type examined as well as specimens from southeastern Alaska — 
Hartman, 1948), and E. cinerea Webster and Benedict (1884) from 
Provincetown, Massachusetts (type in USNM), appear to be 
synonymous with E. longa. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, 1.7-75.5 fms., up to 8 
miles from shore, on bottoms of sandy-mud, stones, and various 
combinations of mud, pebbles, stones, gravel, rocks (12 stations, 40 
specimens). West Coast North America: San Juan Archipelago, 
Washington, Pettibone. West Greenland: Vaigat, Disko Island, 
mud, Bartlett, 1937. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergea, Franz Josef Land, 
Novaya Zemlya. Also Iceland, Norway to English Channel; 
Hudson Bay to Massachusetts; Bering Sea to Mexico; north Japan 
Sea; China. In low water to 518 fathoms. 



]VIARINE POLYCHAETE WORMS — PETTIBONE 235 

Eteoneflava (Fabricius, 1780) 

Figure 27, g 

Nereis flava Fabricius, 1780, p. 299. 

Eieone flava Malmgren, 1865, p. 102, pi. 15, fig. 35.— Bergstrom, 1914, p. 196, 

fig. 74.— Eliason, 1920, p. 27.— Fauvel, 1923, p. 173, fig. 62, e, f.— Augener, 

1928, p. 709.— Annenkova, 1937, p. 158; 1938, p. 145.— Zatsepin, 1948, p. 

113, pi. 29, fig. 12.— Wesenberg-Lund, 1950a, p. 11, pi. 2, fig. 7, c; 1950b, p. 

39; 1951, p. 30. 
Eteone depressa Malmgren, 1865, p. 103, pi. 15, fig. 36. — Southern, 1914, p. 79. 
Eteone sarsi Malmgren, 1867, p. 28, pi. 2, fig. 14. — Webster and Benedict, 1887, 

p. 711. 
Eteone lentigera Malmgren, 1867, p. 29, pi. 2, fig. 13. 

Description. — Length 24-68 mm.; width without setae 1-4 mm. 
(length up to 120 mm. — ^Fauvel, 1923). See key. Color: In life: 
Pinkish flesh and grayish white. 

New records. — ^Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, 8.3-54.6 fms., up to 
7.5 miles from shore, on bottoms of mud, and various combinations 
of mud, stones, gravel, rock, large perforated rocks, shells (10 stations, 
22 specimens). 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Spitsbergen, Novaya Zemlya. Also 
Iceland, Faroes, Norway to English Channel; east coast North Amer- 
ica (Maine); Bering Sea; north Japan Sea. In low water to 471 
fathoms. 

Eteone spetsbergensis Malmgren, 1865 

Figure 27, / 

Eteone spetsbergensis Malmgren, 1865, p. 102, pi. 15, fig. 38.— Th6el, 1879, p. 31, 
pi. 2, figs. 21, 22.— Bergstrom, 1914, p. 202, fig. 77.— Ditlevsen, 1917, p. 66, 
pi. 5, figs. 12, 14, 18.— Eliason, 1920, p. 27.— Augener, 1928, p. 708.— Annen- 
kova, 1937, p. 159; 1938, p. 145.— Hartmau, 1948, p. 20, fig. 5, b.— Berkeley 
and Berkeley, 1948, p. 42.— Zatsepin, 1948, p. 113, pi. 29, fig. 11.— Wesen- 
berg-Lund, 1951, p. 31. 

Description. — Length 17-75 mm., width without setae 0.7-4 mm. 
(length up to 100 mm. — Augener, 1928). See key. Color: In life: 
Pale green ventraUy, reddish brown dorsolateral bands, with dark 
dorsal cirri; orange colored eggs. 

New records. — Arctic Alaska: Off Browerville near Point Barrow; 
off Point Barrow base, 1.7-36 fms., up to A.)i miles from shore, on 
bottoms of sandy-mud and various combinations of mud, gravel, 
rocks, stones, shell (4 stations, 4 specimens). 

Distribution. — Scattered records in the Arctic: Arctic Alaska, Spits- 
bergen, Franz Josef Land, Novaya Zemlya. Also Iceland, Norway 
to Scotland; Bering Sea; western Canada; north Japan Sea. In 1.7- 
40 fathoms. 

261112—54 3 



236 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Eteone sp. larvae 

Larvae of a species of Eteone found in a mass of jelly (off Point 
Barrow base, 1.7 fms., 300 ft, out, on sandy-mud, July 20, 1948); 11 
segments, of which 8 were setigerous; triangular prostomium with 
short antennae and eyes; tentacular segment with short tentacular 
cirri; anal cirri short, spherical. 



Genus Phyllodoce Savigny, 1817 

Subgenus Anaitides Czerniavsky, 1882 
Phyllodoce (Anaitides) groenlandica Oersted, 1842 

Figure 27, d, i 

Phyllodoce groenlandica Oersted, 1842, p. 121 {fide Bergstrom, 1914); 1843, p. 192, 
pi. 2, figs. 19, 21, 22, 29-32.— Malmgren, 1865, p. 96; 1867, p. 21, pi. 2, fig. 
9.— Webster and Benedict, 1884, p. 703; 1887, p. 710.— Murdoch, 1885, p. 
153.— Moore, 1903, p. 428.— Mcintosh, 1908, p. 86, figs.— Ditlevsen, 1917, 
p. 56.— Fauvel, 1923, p. 153, fig. 54, f-i.— Augener, 1928, p. 703.— not 
Treadwell, 1937, p. 28 { — Paranaitis wahlbergi (Malmgren), in USNM). — 
Annenkova, 1937, p. 156.— Friedrich, 1939, p. 122.— Thorson, 1946, p. 52, 
fig. 21.— Gorbunov, 1946, p. 38.— Berkeley and Berkeley, 1948, p. 46, fig. 
66.— Wesenberg-Lund, 1950a, p. 10; 1950b, p. 32; 1951, p. 26. 

Phyllodoce citrina Moore, 1908, p. 328. — Hartman, 1948, p. 19 (part; includes 
P. groenlandica and P. mucosa, in USNM; not P. citrina Malmgren, 1865). 

Anaitides groenlandica Bergstrom, 1914, p. 141, fig. 42. — Southern, 1914, p. 68. — 
EUason, 1920, p. 24 (part) .— Chamberiin, 1920, p. 11.— Hartman, 1944a, 
pp. 335, 338; 1948, p. 19.— Zatsepin, 1948, p. Ill, pi. 29, fig. 2. 

Description. — Length 16-285 mm., width without setae 1-9 mm. 
(length up to 450 mm. — Ditlevsen, 1917). Segments very numerous. 
Body elongate, large, robust, linear, flattened, attenuate posteriorly. 
Prostomium cordiform, notched posteriorly with an occipital tubercle 
in the notch, with foiu- short subulate frontal antennae, with smaU, 
paired, lateral, retractile, nuchal knobs just anterior to the first pair 
of tentacular cirri. Tentacular cirri unequal; two pairs short, two 
pairs longer, extending to segments 9-11. Dorsal cirri of median 
region up to two times longer than wide, subquadrangular, with distal 
ends truncate; ventral cirri oval, with asymmetrical acuminate tip. 
Anterior part of extended proboscis transversely rugose, crowned 
mth 17 papillae; basal part with 12 longitudinal rows of smaU, oval 
papillae (6 rows on each side), with 10-20 papillae per row. Color: 
In life and in alcohol: Body with bluish iridescence, usually heavily 
pigmented dark greenish brown or deep reddish blue, with dorsal and 
ventral cirri tan to dark brown; on some, body not so darkly pig- 
mented, with cirri only partly pigmented. 



MARINE POLYCHAETE WORMS — PETTIBONE 



237 




Figure 27. — Phyllodocidae: a, Eteone, dorsal view anterior end; b, My slides borealts, 
dorsal view anterior end; c, Eumida minuta, dorsal view anterior end; d, Phyllodoce 
groenlandica, dorsal view anterior end; e, Eteone barhata, parapodium; /, Eteone spets- 
bergensis, parapodium; g, Eteone flava, parapodium; h, Eteone longa, parapodium; i, 
Phyllodoce groenlandica, parapodium. (For explanation of symbols, see p. 210.) 



238 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Remarks. — Some smaller specimens were at first placed under P. 
citrina Malmgren, since the papillae on the proboscis were fewer than 
normal, four rows on each side, with 3-5 papillae per row. An inter- 
mediate-sized specimen from Canoe Bay, Alaska (Hartman, 1948, as 
P. citrina) has five rows on each side, with as many as 6 papillae per 
row. It appears that the papillae are added on gradually and that 
only in the fully developed individual is the full complement of 6 
rows of papillae on each side developed ; the number per row is varia- 
ble, ranging from 10 (^vith some spaces as if some had dropped off) 
to 20. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow; Point Barrow base, washed ashore; off Point Barrow 
base, up to 12.1 miles from shore, 1.7-123.5 fms., on bottoms of mud, 
sandy-mud, mass of worm tubes, and various combinations of mud, 
gravel, stones, pebbles, rocks, large perforated rocks, shells; in screen 
trap on bottom (23 stations, 74 specimens). West Greenland: 
Vaigat, Disko Island, mud, Bartlett, 1937. East Coast North 
America: Off Labrador, 70 fms., Blue Dolphin Expedition, 1950; off 
Maine, Massachusetts, intertidal to 42 fms., U. S. Fish Commission. 
West Coast North America: Washington Sound, Pettibone. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Spitsbergen, Franz 
Josef Land, Barents Sea, Novaya Zemlya, Kara Sea; Also Iceland, 
Faroes, Norway to English Channel; Hudson Bay to Massachusetts; 
Bering Sea to Washington; north Japan Sea. In low water to 800 
fathoms. 

Genus Eumida Malmgren, 1865 
Eumida minuta (Ditlevsen, 1917) 

Figure 27, c 

Eulalia minuta Ditlevsen, 1917, p. 56, pi. 4, figs. 10, 12, 14. — ? Gorbunov, 1946, 

p. 38. 
Eulalia arctica Aunenkova, 1946, pp. 185, 187, fig. 1, a-c. 

Description. — Length 1-8 mm.; width without setae 0.3-0.8 mm. 
Segments few (12-36). Body very small, slightly tapered anteriorly 
and posteriorly, flattened dorsoventrally. Prostomium semiglobular 
to bluntly conical, with two large eyes near posterior border, with 
antennae rather long, slender, filamentous, almost as long as pro- 
stomium. First tentacular achaetous segment not distinct dorsally, 
with first pair tentacular cirri lateral to prostomium at same level as 
the eyes (thus placed under Eumida and not Eulalia Savigny); two 
pairs tentacular cirri on second segment (first setigerous); fourth 
tentacular cirri on third segment; tentacular cirri enlarged basally, 



MARINE POLYCHAETE WORMS — PETTIBONE 239 

with slender filamentous tips, the latter about as long as the enlarged 
basal part except for the ventral tentacular cirri on the second segment 
which are much shorter. Dorsal cirri oval or bluntly conical. 
Ventral cirri elongate oval. Anal cirri short, oval, 1.5 to 2 times 
longer than wide. Color: In alcohol: Greenish tan to brown. In 
life: Flesh ventrally, green and flesh dorsally, with salmon-colored 
eggs showing through. 

New records. — Arctic Alaska: Off Point Barrow base, 16.7-75.5 
fms., up to 8 miles from shore, on various combinations of gravel, 
small stones, rocks, bryozoans, hydroids, Saxicava shells (6 stations, 
46 specimens). 

Distribution. — Few scattered records in the Arctic: Siberian and 
Alaskan Arctic, Davis Strait. In 16.7-75.5 fathoms. 

Family Hesionidae 

Prostomium usually with four eyes, two or three antennae, two 
biarticulate palps (fig. 28, a). Two to eight pairs tentacular cirri on 
one to four more or less distinct achaetous tentacular segments. 
Parapodia biramous or subbiramous; notopodia may be greatly re- 
duced, with notosetae simple or lacking; neurosetae compound (fig. 
28, b). Dorsal cirri long, more or less distinctly articulated; ventral 
cirri shorter; two long anal cirri. Proboscis cylindrical, eversible, 
with marginal papiUae, with or without horny jaws. 

Represented by a single species from Point Barrow. (The speci- 
mens from Point Barrow, referred to Castalia multipapillata Theel by 
Murdoch (1885), do not agree with this species nor with any other 
species of hesionid; the parapodia are biramous, both lobes being 
pointed, the notopodia are well developed, with capillary setae; 
neurosetae compound, with delicate tips; the proboscis has numerous 
small papillae; tentacular cirri four pairs; prostomium with two 
pairs eyes, pair biarticulate palps, antennae ? (missing or absent); 
the specimens are very small and in poor condition.) 

Genus Castalia Savigny, 1820; emend. Fauvel, 1923 
Castalia aphroditoides (Fabricius, 1780) 

Figure 28, a, h 

Nereis aphroditoides Fabricius, 1780, p. 296. 

Castalia ardica Malmgren, 1867, p. 32. — Annenkova, 1931, p. 203. 

Castalia fabricii Malmgren, 1867, p. 32.— Th6el, 1879, p. 37, pi. 3, figs. 36, 37.— 

Berkeley and Berkeley, 1943, p. 130. 
Castalia aphroditoides Wir6n, 1883, p. 401.— Augener, 1913, p. 260; 1928, p. 715.— 

Zatsepin, 1948, p. 114.— Wesenberg-Lund, 1950a, p. 13, pi. 3, fig. 14; 19o0b, 

p. 44. 
Psammate aphroditoides Chamberlin, 1920, p. 13. 



240 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Description. — Length 2.5-18 mm., width without setae 0.7-2.5 mm. 
Segments 25-43. Body cylindrical. Prostomium rectangular, wider 
than long, widest anteriorly, notched posteriorly, with two pairs 
eyes in trapezoidal arrangement, anterior pair larger; palps on an- 
terolateral borders of prostomium; two filiform antennae medial to 
palps (without unpaired antenna); antennae subequal, about same 
length as prostomium. SLx pairs tentacular cirri on three more or less 
distinct achaetous segments; tentacular cirri long, filiform, articulated, 
with prominent cirrophores. Without distinct notopodia (acicula in 
cirrophores of dorsal cirri only); without notosetae. Neuropodia 
well developed, with three diverging conical distal lobes : Lip anterior 
to setae with median and dorsal conical lobes; lip posterior to setae 
with rounded median and conical ventral lobes; neurosetae compound. 
Dorsal and anal cirri long, articulated. Proboscis barrel-shaped with 
10 soft papillae around opening (four dorsal to lateral grooves, three 
each on two ventrolateral folds), with a median ventral notch, with 
two thickened ventrolateral ridges (the so-called jaws) lateral to the 
ventral notch. Color: In alcohol: Without color, or slightly greenish 
dorsally (especially in anterior part), or brownish. In life: Orange- 
flesh, with greenish eggs inside, with red eyes. 

Remarks. — Two young specimens found in plankton tow beneath 
ice February 28, 1950. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms., stony-mud; off Point Barrow base, 21-75.5 
fms., up to 8 miles from shore, on bottoms of mud, stones, and various 
combinations of mud, stones, gravel, rocks, and from breaking apart 
fohaceous brj^ozoans (10 stations, 28 specimens). Northwest 
Greenland: Walrus grounds, Murchison Sound, Bartlett, 1938. 
East Coast North America: Off Labrador, 45 fms.. Blue Dolphin 
Expedition, 1949. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Franz Josef Land, 
Novaya Zemlya, Kara Sea. Also Bering Sea; Denmark; Hudson Bay, 
Labrador. In 2-75.5 fathoms. 

Family Syllidae 

Body of small size. Prostomium with four eyes (sometimes six), 
two palps (may be reduced and fused), three antennae (figs. 28, 29). 
Tentacular segment achaetous, with one or two pairs tentacular cirri. 
Parapodia uniramous, with dorsal and ventral cirri (latter may be 
absent). Two anal cirri. Setae mostly compound, sometimes with 
additional simple setae. Proboscis eversible, armed or not with one 



MARINE POLYCHAETE WORMS — PETTIBONE 241 

to several chitinous teeth. Reproduction direct (epigamy) or by 
stolons, sometimes having sexual dimorphism; with swimming capillary 
setae at maturity. 

Represented by 6 genera and 11 species. 



Key to the genera of Syllidae from Point Barrow 

1. Without ventral cirri. Palps reduced, united, turned under on ventral side 

of prostomium (except for forked palps of male stolon; fig. 29, a). Antennae 
and dorsal cirri smooth or faintly annulated. Reproduction by stolons; 
stem forms produce by transverse fission, singly or in chains, sexually 

dimorphic male and female forms with swimming setae Autolytus — 2 

With ventral cirri (fig. 28, d) . Palps better developed, free or fused 4 

2. Prostomium normal, with 4 dorsal eyes in trapezoidal arrangement, with 3 

antennae, short fused palps; 2 pairs tentacular cirri (fig. 29, a). Without 
swimming setae. Pharynx long, more or less sinuous, usually with a crown 

of teeth Stem form or stock of Autolytus (p. 242) 

Prostomium abnormal, with 4 large eyes (usually 2 dorsolateral and 2 ventro- 
lateral) ; 2-3 pairs tentacular cirri (fig. 29, c-f) . Body divided into an anterior 
unmodified region, a middle region with long swimming setae and well- 
developed, paddlelike parapodia, and with or without a posterior unmodified 
region. Without pharynx Sexual stolons of Autolytus — 3 

3. Prostomium with 3 antennae (very long median one and pair of small frontal 

ones) and pair of anterior bifurcate palps; 1 pair tentacular cirri very long, 
similar to median antennae (these may be the dorsal cirri of the first setiger 
or of a small, knoblike achaetous lobe; fig. 29, e,f). Testes and sperm con- 
fined to anterior unmodified region. 

Male form of Autolytus (Polybostrichus) (p. 242) 
Prostomium with 3 subequal antennae, without palps; none of tentacular 
cirri especially elongate (fig. 29, c, d). Most of body filled with eggs; eggs 
carried in large sac on ventral surface (gestation). 

Female form of Autolytus (Sacconereis) (p. 242) 

4. Dorsal cirri distinctly moniliform throughout body (fig. 28, c) . Palps separate 

for their entire length or fused basally. Reproduction direct or by single 
stolons. Sexual stolons (Chaetosyllis) : Small prostomium, with 2 pairs 
eyes, pair short moniliform antennae; without tentacular cirri; 1 anterior 
unmodified segment, a middle region with long swimming setae, and few 

unmodified posterior segments Syllis (p. 252) 

Dorsal cirri smooth or indistinctly articulate (may be more or less articulate 
anteriorly) . Palps fused at base only or throughout .__ 5 

5. Palps fused for nearly their entire length (fig. 28, k-m). Body very small. 

Antennae and dorsal cirri short, not moniliform. One pair tentacular cirri 
(may be rudimentary). Reproduction generally direct, with swimming 
setae at maturity; eggs and larvae fixed to dorsal or ventral surface of 

female 6 

Palps fused at base only (fig. 28, g). Body larger. Antennae and dorsal 
cirri longer, smooth (antennae and anterior dorsal cirri may be indistinctly 
articulate, especially distally). Two pairs tentacular cirri. Reproduction 
direct, with swimming setae at maturity 7 



242 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

6. Antennae and dorsal cirri swollen at base, tapering to narrow tip (fig. 28, m) . 

Body and parapodia usually covered with adhesive papillae and fine, granular 

material Sphaerosyllis (p. 255) 

Antennae and dorsal cirri clavate or conical (fig. 28, k, I). Bodj' and parapodia 
not covered with papillae Exogone (p. 257) 

7. Compound neurosetae with distal blades all rather short (fig. 28, i). Pro- 

boscis with a distal and proximal row of soft papillae, with distal circular 

chitinous margin denticled (fig. 28, g, h) Eusyllis (p. 259) 

Compound neurosetae with some distal blades elongate (fig. 28, j). Proboscis 
with distal row of soft papillae only, with distal circular chitinous margin 
smooth Pionosyllis (p. 262) 



Genus Aiitolytus Grube, 1850 

In working over the rather numerous specimens of Autolytus in 
the collections from Point Barrow, several things were revealed which 
throw light on the confusion in this group caused by the formation of 
sexually dimorphic stolons. For correct description of any species, 
one should have the stem form, the sexual buds in the process of 
formation including the mature sexual buds attached to the stem 
form, and the mature male and female stolons separated from the 
stem. Very often, however, stem forms and the male and female 
stolons of one species have been described and given different names, 
while sexual stolons and stem forms of different species have been 
given the same name. A. fallax Malmgren, described originally from 
specimens from Spitsbergen, is of particular interest. Malmgren 
described and figured the stem form -with the head of a male stolon 
forming between se tigers 13 and 14 and indicated that the sexual 
stolons were not known. A. prismaticus (Fabricius), described 
originally from specimens from Greenland, has a characteristic color 
pattern of three dark longitudinal bands (median and lateral at the 
level of the bases of the dorsal cirri). The sexual stolons have six 
unmodified setigers anterior^toHhe setigers' with swimming setae; the 
stolons show the characteristic color pattern of the stem form. In 
the absence of color, it would be difficult to distinguish A. prismaticus 
from certain other species of Autolytus. It apparently has been the 
usual practice to identify any sexual stolon of unknown connections 
that has six anterior unmodified setigers as A. prismaticus, and some- 
times to identify the male stolons as Polyhostrichus longosetosus 
Oersted or A. longisetosus (Oersted). However, as found in the Point 
Barrow material, A. fallax is the most common species of Autolytus 
and it was found along \vith A. prismaticus and A. alexandri. The 
majority of the specimens of A. Jallax were in the process of stolon 
formation, the head of the stolon being formed between setigers 13 



MARINE POLYCHAETE WORMS — PETTIBONE 



243 




Figure 28. — Hesionidae: a, Castalia aphroditoides, dorsal view anterior end; b, same, 
parapodium, posterior view. Syllidae: c, Syllis fasciata, dorsal view anterior end; d, 
same, parapodium; e, same, neuroseta; /, Syllis cornuia, neurosetae, with short (1) and 
long (2) blades; g, Eusyllis blomstrandi, dorsal view anterior end, proboscis partially 
extended; h, same, frontal view extended proboscis showing distal row of papillae and 
denticled margin; t, same, neuroseta; /, Pionosyllis compacta, neurosetae, lower one (1) 
with shorter blade, upper one (2) with longer blade; k, Exogoru dispar, dorsal view anterior 
end; /, Exogone naidina, dorsal view anterior end; m, Sphaerosyllis erinaceus, dorsal view 
anterior end. (For explanation of symbols, see p. 210.) 



244 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

and 14. Although no mature sexual buds were found attached to the 
stem form, the rudiments of the notopodia with the developing swim- 
ming setae could be seen, beginning on setiger 7. Male and female 
stolons with six anterior unmodified segments were found. Of these, 
two female stolons showed the characteristic banding of A. prismaticus 
(this banding was noted on aU the developing sexual buds still attached 
to the stem forms of this species). The others did not show this 
banding and are considered to be the sexual stolons of A. jallax. A. 
Jallax, then, along with A. prismaticus, has six anterior, unmodified 
setigers and manj^ of the records of A. prismaticus in the literature 
will no doubt prove to be A. jallax. 

The stem form of A. fallax has been referred to A. prolijer (Miiller) 
by Augener (1913, 1928) and Wesenberg-Lund (1947), thus the stem 
form of A. fallax has been confused wdth A. prolifer and sexual forms 
with A. prismaticus. 

There appears to be a correlation in the place of formation of the 
sexual stolons in the stem form with the number of anterior, unmodi- 
fied setigers and the number of tentacular cirri in the sexual stolons. 
At least for a group of species, including A. prismaticus, A. fallax, A. 
cornutus A. Agassiz, and A. ornatus (Verrill, 1873), the stem forms 
produce sexual stolons singly, with the head of the stolon forming 
between setigers 13 and 14. The stolon, then, consists of the segments 
of the stem posterior to setiger 13 (differing from A. prolifer where 
the sexual stolons are proliferated more posteriorly, often in chains of 
2 to 8). The posterior part develops gradually into the male or 
female sexual stolon and breaks off when mature. The anterior 13 
setigers of the stem regenerates another posterior end. The sexual 
stolons of these four species have six mimodified setigers anterior to 
the swimming setae and usually a more or less developed immodified 
posterior region. The stolons have three pairs of tentacular cirri on 
the first two achaetous tentacular segments; the first two pairs on the 
first segment and the third pair on the second segment. The upper 
pair of tentacular cirri on the first segment are similar to the dorsal 
cirri. The lower pair may be very short, about the length of the 
prostomium. The third pair of tentacular cirri in the female are 
similar to the dorsal cirri; in the male they are very long, similar to 
the median antenna. In addition, there are a pair of small achaetous 
lobes. In contrast, in A. prolifer the sexual stolons have three ante- 
rior unmodified setigers (sometimes two or four) and a poorly devel- 
oped posterior region; there is a single tentacular achaetous segment 
with two pairs of tentacular cirri (in the female both the upper and 
lower pairs are short; in the male the lower one is similar to the dorsal 
cirri and the upper one is very long, similar to the median antenna). 
In A. alexandri the sexual stolons have 14 (sometimes 13) anterior 



MARINE POLYCHAETE WORMS — PETTIBONE 245 

unmodified setigers and a well-developed posterior region; there is a 
single achaetous tentacular segment with two pairs of tentacular cirri; 
the first setigerous segment in the female is similar to the following 
segment, but in the male it has a very long pair of dorsal cirri similar 
to the median antenna. Thus the male stolons of the different species 
of Autolytus agree in having a very long pair of anterior cirri similar 
to the median antenna, but they may be developed as the upper pair 
of tentacular cirri of a single tentacular segment as in ^. prolifer, as 
the dorsal cirri of the first setigerous segment as in ^. alexandri, or as 
the third pair of tentacular cirri on the second tentacular segment 
just dorsal to a pair of achaetous lobes as in A. prismaticus. 

The number of setigers in the anterior, unmodified region of the 
sexual stolons appears to be a good taxonomic character for most 
species. That the number is somewhat variable in A. prolifer (usually 
three, sometimes two and four) does not mean that it is of no signifi- 
cance in other forms. Thus the specimens of Autolytus with six 
anterior unmodified setigers, which Dales (1951) referred to A. prolifer, 
are no doubt another species. The description agrees well with that 
of A. cornutus. 

All three species of Autolytus from Point Barrow have the body 
linear elongate, attenuated posteriorly, flattened ventrally, arched 
dorsally. Prostomium with four eyes in trapezoidal arrangement, 
with palps fused, turned ventrally, exceeding the prostomium only 
sUghtly. Neuropodia with composite setae with blades short, biden- 
tate, and usually with a simple bayonette seta. 

Key to the species of Autolytus from Point Barrow 

1. Stem Form: Dorsal cirri irregular in length, at least some longer than body 
width, pigmented with reddish globules; body whitish, brownish, or trans- 
versely banded with reddish globules, 2 narrow bands per segment; with 
nuchal epaulettes on first 2 (to 4) setigers; sexual buds formed singly (?), 
the heads forming posterior to setiger 14. Sexual Stolons: 14 setigers 
anterior to swimming setae; nuchal epaulettes on first 3 setigers; 2 pairs 
tentacular cirri, the upper one longer; in male, first pair dorsal cirri on first 
setiger very long, similar to median antenna; female stolons with 2-lobed 

egg sac A. alexandri 

Stem Form: Dorsal cirri, except anterior 2 pairs, shorter than body width, 
not extending beyond or only slightly be3'ond the setae, subequal; sexual 
buds formed singly, the heads forming between setigers 13 and 14. Sexual 
Stolons: 6 setigers anterior to swimming setae; 3 pairs tentacular cirri; in 
male, first pair similar to dorsal cirri, with second short pair at its ventral 
base, third pair very long and stout (similar to median antenna) with small 
achaetous lobe at its ventral base (fig. 29, e, f) ; in the female, similar except 
both the first and third pairs are similar to the following dorsal cirri (fig. 
29, c, d); female stolons with 1-lobed egg sac 2 



246 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

2. Stem Form: Usually with 2 dorsolateral longitudinal black to brown bands 
just dorsal to bases of dorsal cirri (may be faint or absent); with short 
semicircular epaulettes on tentacular segment only. Sexual Stolons: 
Colorless or female stolon dusk}'^, especially posteriorly; male stolon rusty 
brown, darkest on anterior region, especially laterally; epaulettes on 

tentacular segment only (fig. 29, c-f) A. fallax 

Stem Form: With 3 longitudinal black bands, 1 medium and 2 lateral at same 
level as bases of dorsal cirri; medium band continuous throughout length of 
body, lateral ones may fade out in anterior fourth or be lacking; with semi- 
circular epaulettes on tentacular segment and part of first setiger (fig. 29, a). 
Sexual Stolons: With 3 longitudinal pigmented bands, median one con- 
tinuous throughout most of body, lateral ones along bases of dorsal cirri 
in anterior unmodified region; with epaulettes on tentacular segment and 
part of first setiger A. prisxnaticus 

Autolytus alexandri Malmgren, 1867 

Autolytus alexandri Malmgren, 1867, p. 37, pi. 7, fig. 39. — Verrill, 1881, p. 292, 

pi. 12, fig. 8.— Chamberlin, 1920, p. 12.— Friedrich, 1939, p. 123.— Hartman, 

1942a, p. 13; 1944a, p. 338, pi. 13, fig. 11 (not pi. 13, fig. 2); 1945, p. 17, 

pi. 2, fig. 11. 
Stephanosyllis pida Verrill, 1874a, pp. 43, 132, pi. 4, fig. 6; 1874b, pp. 361, 362, 

pi. 4, fig. 1 (not Proceraea picta Ehlers, 1864). 
Stephanosyllis ornata Verrill, 1874a, p. 132 (not Proceraea ornata Verrill, 1873). 
Proceraea (Stephanosyllis) ornata Webster and Benedict, 1887, p. 724 (not 

Proceraea ornata Verrill, 1873). 
Autolytus verrilli Marenzeller, 1892, p. 416, pi. 19, fig. 4. — Augener, 1928, p. 726. — 

Wesenberg-Lund, 1947, p. 33, figs. 14, 15; 1950a, p. 18; 1950b, p. 52; 1951, 

p. 52. 

Description. — Stem form: Length 4-12 mm., width without setae 
0.6-0.8 mm., up to 35-64 segments. Antennae rather short, thick, 
subequal or median one up to twice the length of the lateral; tentac- 
ular cirri equal to or shorter than antennae, upper pair longer; 
first pair dorsal cirri as long as or longer than median antenna; rest 
of dorsal cirri rather irregular, unequal, at least some longer than body 
width, with cirrophores prominent. Distinct epaulettes on first two 
setigerous segments (or to end of setiger 4 — Hartman, 1945). None 
showed sexual bud formation (sexual bud is formed between segments 
25 and 26 — Hartman, 1945). Color: In alcohol: Colorless or trans- 
versely banded with reddish to brownish granules, may be two bands 
per segment. 

Female stolon (Sacconereis) : A single specimen of 72 setigers, 
16 mm. long, 1.2 mm. wide without setae, 3.5 mm. wide with swimming 
setae, consisting of 14 anterior, 26 middle, and 32 posterior setigers. 
Prostomium rounded, with four large dorsal eyes, three antennae 
rather short, subequal; two pairs tentacular cirri of nearly same 
length as antennae; dorsal cirri of anterior part of unequal length but 
all much longer than body width. Thick epaulettes extending on 
first three setigers. The stolon, with swimming setae and fiUed with 



MARINE POLYCHAETE WORMS — PETTIBONE 247 

eggs, was dredged in 27 fathoms February 18, 1950. (A 2-lobed 
egg mass is carried ventrally — Wesenberg-Lund, 1947.) 

Male stolon (Polyhostrichus) : No specimens taken from Point 
Barrow. A specimen taken in plankton at Portage Bay, Alaska, was 
12 mm. long, 1.5 mm. wide without setae, 4 mm. wide including 
swimming setae, with 61 setigers consisting of 14 anterior, 36 median, 
and 11 posterior segments. Prostomium similar to A. fallax; two 
pairs tentacular cirri, upper pair longer. First pair dorsal cirri on 
first setiger very long, similar to median antenna (achaetous — ^Wesen- 
berg-Lund, 1947); dorsal cirri in anterior region long, unequal; in 
middle region short, about one-third length of elongated parapodial 
lobes. Large epaulettes on first three setigers. 

New records. — Arctic Alaska: Off Point Barrow base, 16.7-123.5 
fms., up to 15 mUes from shore, on bottoms of stones, worm tubes, and 
various combinations of mud, pebbles, gravel, stones, rocks, large 
perforated rocks, with bryozoans, hydroids, shells, and worm tubes 
(17 stations, 53 specimens). West Coast North America: Albatross 
surface station, Portage Bay, Alaska; San Juan Channel, Washington 
Sound, 25 fms., Pettibone, 1940. East Coast North America: Off 
Labrador, 30-40 fms.. Blue Dolphin Expedition, 1949; Woods Hole 
region, Massachusetts, Pettibone, 1950. 

Distribution. — Scattered records in the Arctic: Arctic Alaska, 
Greenland, Spitsbergen, Franz Josef Land, Barents Sea. Also Iceland; 
Labrador to North Carolina; Bering Sea to Washington. In low 
water to 123.5 fathoms; sexual forms at surface. 

Autolytus fallax Malmgren, 1867 

Figure 29, c-f 

? Polyhostrichus longosetosus Oersted, 1843, p. 183, pi. 5, figs. 62, 67, 71. — Tread- 

weU, 1937, p. 28. 
Autolytus fallax Malmgren, 1867, p. 33, pi. 6, fig. 41.— Ditlevsen, 1929, p. 17. 
? Autolytus longisetosus Malmgren, 1867, p. 34, pi. 7, fig. 38. 
Autolytus prolifer Augener, 1913, p. 258; 1928, p. 724. — Wesenberg-Lund, 1947, 

p. 19, figs. 8, 9 (part; not Nereis prolifera Miiller, 1788). 
Autolytus prismaticus Wesenberg-Lund, 1947, p. 24 (part; not Nereis prismatica 

Fabricius, 1780). 

Description. — Stem form: Length 3-18 mm., width without setae 
0.3-0.7 mm,, up to 45-78 segments. Median antenna and first pair 
dorsal cirri very long; lateral antennae and upper tentacular cirri 
about half as loDg; lower pair tentacular cirri and second pair dorsal 
cirri shorter, about twice the length of the following cirri ; rest of dorsal 
cirri short, less than half the body width and extending only slightly 
beyond the setae, subequal. Short epaulettes (a pair of shallow, semi- 
circular raised areas) on tentacular segment only. Of 68 specimens of 
the stem form, 45 showed bud formation of the sexual stolons, the 



248 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

head of the bud forming between setigers 13 and 14; 6 were broken 
after setiger 13, 5 consisted of 13 setigers plus a definite posterior 
growing zone, and 12 consisted of 36 setigers or less and showed no 
sign of bud formation. Of the 45 specimens showing bud formation, 
18 were young buds (sex?) showing four eyes and rudiments of head 
appendages; 17 were developing male stolons as indicated especially 
by the developing forked palps (at first, one part of fork about one- 
half the length of the other), large median antenna, and large dorsal 
tentacular cirri; the male buds consisted of a variable number of 
setigers (14-26 plus a definite growing zone, or 32-42 and tapered 
gradually to an attenuated posterior end); 10 were developing female 
buds as indicated by the developing three subequal antennae and the 
absence of forked palps; the female buds consisted of 40-65 setigers. 
Color: In alcohol: Two dorsolateral, longitudinal dark bands just 
dorsal to bases of dorsal cirri (almost all specimens showed at least 
faint bands). In life: Anterior part pale yellowish with dark dorso- 
lateral bands, posterior part orange. Tubes : Thin, tough, transparent, 
on hydroids as Lajoeina maxima Levinsen; either on the surface or 
surrounded by the sessile hydrothecae of the hydroid colony. 

Female stolon (Sacconereis) : Length 5-15 mm., width without 
setae 0.5-1 mm., width including swimming setae 3.5-4 mm., up to 
26-51 segm^ents. Body widest in median region, tapering slightly 
anteriorly and more so posteriorly, forming an attenuated tail region; 
divided into three regions composed of 6 anterior unmodified setigers, 
13-18 modified setigers with long, iridescent swimming setae, and 
3-27 posterior unmodified setigers. Prostomium short, broad, one 
pair eyes dorsolateral, one pair larger ones ventrolateral, both with 
lenses; antennae long, stout, subequal or median one slightly longer 
than lateral pair. Upper two pairs tentacular cirri similar to dorsal 
chri of following segments, with a third pair of short, ventral, tentacu- 
lar cirri (about length of prostomium) and a pair of small, ventral, 
achaetous lobes. Dorsal cirri about as long as body width. Pair 
of shallow semicircular epaulettes on tentacular segment only. Pyri- 
form-shaped egg sac carried on ventral surface in region of setigers 
10-18, the egg mass flattened or slightly concave on side toward body, 
the narrower end of lobe anterior, the eggs orange-colored (in life). 
Color: In alcohol: Colorless, transparent, may be dusky posteriorly 
or somewhat dusky throughout. Two female stolons with eggs in- 
side body dredged in 75 fms. October 11, 1949; females with egg sacs 
taken from vertical plankton hauls in 13 fathoms March 29, April 
15, May 17, 1950; female with egg sac containing developing embryos 
dredged in 5 fathoms January 25, 1950. 

Male stolons (Polybostrichus) : Length 7-9 mm., width without 
setae 0.8-1 mm., width including swimming setae 2 mm., 43-66 seg- 



MARINE POLYCHAETE WORMS — PETTIBONE 249 

ments, with body regions of 6 anterior, 27-30 middle, 9-10 posterior 
setigers, tapering gradually posteriorly. Prostomium short, wide, 
with a pair of eyes dorsolateral, a larger pan- ventrolateral, all with 
lenses; palps bifurcated about basal third, the two branches subequal; 
very long, stout median antenna and very small lateral antennae 
anterior and medial to dorsal pair of eyes. Three pairs tentacular 
cirri with anterior dorsal pair similar to dorsal cirri, anterior ventral 
pair shorter than prostomial length, posterior dorsal pair very large, 
similar to median antenna, with a pair of achaetous knobs at their 
ventral bases. Dorsal cirri of anterior setigers about as long as body 
width, those of modified region about as long as the elongated, paddle- 
like parapodial lobes, those of posterior, taillike region short, digitiform. 
Shallow semicircular epaulettes on tentacular segment only. Color: 
In alcohol: Rusty brown, darkest on anterior part or colorless except 
anterior part. In life : Yellowish green. Taken in plankton, near shore, 
September 10, 1949; dredged in 27 fathoms February 18, 1950. 

Remarks. — As referred to under the systematic discussion of 
Autolytus, the stem form of A. jallax has been incorrectly referred to 
A.'proliferhj KugmQv (1913, 1928) and Wesenberg-Lund (1947, 1950). 
The two may be distinguished by the different positions of the sexual 
buds, the greater length of the dorsal cirri in A. prolifer, and the color 
markings. The sexual stolons of A. jallax have been confused with 
A. prismaticus (see discussion above). 

New records.— -Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 5-6.6 fms., mud, in tubes associated with hydroid 
Lafoeina maxima Levinsen; Point Barrow base, washed ashore, on 
same hydroid; off Point Barrow base, 5-75.5 fms., up to 8 miles 
from shore, on bottoms of mud, stones, and various combinations of 
mud, gravel, stones, rocks, large perforated rocks, with bryozoans, 
hydroids including Lafoeina maxima, and shells (16 stations, 85 
specimens); sexual stages from plankton near shore to 75.5 fms. (7 
stations, 22 specimens). Cape Smyth, Point Barrow Expedition, 
winter, 1883 (1 male, 2 females with egg sacs). East Coast North 
America: Off Labrador, 25-60 fms., Blue Dolphin Expeditions, 1949, 
1951. 

Distribution. — Widely distributed in the Arctic: Alaskan and 
Canadian Arctic, Greenland, Spitsbergen, Franz Josef Land, Novaya 
Zemlya. Also Faroes; Labrador. In low water to 75.5 fathoms; 
sexual stolons in plankton. 

Autolytus prismaticus (Fabricius, 1780) 

Figure 29, a, b 

Nereis prismatica ? MuUer, 1776, p. 218.— Fabricius, 1780, p. 302; 1799, p. 177, 

pi. 4, figs. 17-20. 
? Polybostrichus longosetosus Oersted, 1843, p. 183, pi. 5, figs. 62, 67, 71. 



250 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

? Autolytus longisetosus Malmgren, 1867, p. 34, pi. 7, fig. 38. — Verrill, 1881, p. 

292, pi. 12, fig. 10.— Moore, 1902, p. 274; 1909b, p. 134. 
Autolytus incertus Malmgren, 1867, p. 35, pi. 6, fig. 40. 
Proceraea gracilis Verrill, 1874a, pp. 43, 132, pi. 5, fig. 1; 1874b, pp. 361, 362, 

370, 379, pi. 3, fig. 2.— Webster and Benedict, 1887, p. 723. 
Autolytus prismaticus Marenzeller, 1892, p. 420. — Chamberlin, 1920, p. 12. — 

Augener, 1928, p. 725.— Annenkova, 1934, p. 322; 1938, p. 156.— Not 

Berkeley and Berkeley, 1938, p. 48; 1948, p. 68.— Gorbunov, 1946, p. 38.— 

Wesenberg-Lund, 1947, p. 24, figs. 10-12 (part). — ? Hartman and Reish, 

1950, p. 13. 
Autolytus trilineatv^ Berkeley and Berkeley, 1945, p. 318, fig. 1, a-b; 1948, p. 

69, fig. 100. 

Description.- — ^Stem form: Length 5-24 mm., mdth without setae 
0.7-1 mm., up to 52-92 segments. Proportion of anterior appendages 
and dorsal cirri similar to A. fallax. Distinct epaulettes on tentacular 
segment and usually extending also on at least part of first setigerous 
segment. Of 7 specimens, 2 were without signs of bud formation, 
although the body was constricted more than normally between 
setigers 13 and 14; 4 showed sexual bud formation with the head 
developing between setigers 13 and 14; and 1 had 13 setigers mth a 
newly regenerating posterior end. Color: In alcohol: Creamy white 
with three conspicuous, longitudinal, black or dark bands, the median 
one broader, less dense, continuous throughout body, the dorsolateral 
ones narrower, darker, at level of bases of dorsal cirri, often confined 
to anterior fourth or half of body. 

Female stolon (Sacconereis) : A specimen of 58 setigers, 9 mm. long, 
1 mm. wide without setae, 3 mm. wide with swimming setae, with 
body regions of 6 anterior, 18 middle, and 34 posterior setigers. 
Epaulettes somewhat triangular, extending on tentacular and first 
setigerous segments. Otherwise SiS in A. fallax. Color: In alcohol: 
Shows the same characteristic pigmentation as the stem form, three 
longitudinal, black pigmented bands. Two female stolons with 
swimming setae and eggs massed inside body dredged in 5 and 6 
fathoms August 6, 30, 1948. Male stolons {Polyhosirichus) : no 
specimens taken. 

Remarks. — Polyhosirichus longosetosus Oersted, the male stolon 
(this might well be the male stolon of A. fallax, see systematic dis- 
cussion above), Autolytus incertus Malmgren, the female stolon, and 
Proceraea gracilis Verrill, the stem form, have been referred previously 
by Marenzeller (1892) to A. prismaticus. A. trilineatus Berkeley and 
Berkeley is herein referred to A. prismaticus] the description, including 
the characteristic three longitudinal bands, is in agreement. The 
record of A. prismaticus by the Berkeleys (1938, 1948) is doubted 
since the stem form agrees with A. cornutus or A. prolifer but not A. 
prismaticus; the sexual forms agree with A. cornutus or A. prismaticus 
in the number of six anterior unmodified setigers; they agree with A. 



MARINE POLYCHAETE WORMS — ^PETTIBONE 



251 




^frkn ,p 



tCi"' 





Figure 29. — Syllidae: a, Autolytus prismaticus, stem form, dorsal view anterior end; h 
same, parapodium; c, Autolytus fallax, female (Sacconereis) form, dorsal view anterior 
end; d, same, ventral view anterior end; e, Autolytus fallax, male (Polybostrichus) form, 
dorsal view anterior end; /, same, ventral view anterior end. (For explanation of symbols, 
see p. 210.) 



261112—54- 



252 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

comutus as to size, about one-third the size of the sexual stage of A. 
prismaticus, and in the absence of semicircular epaulettes (specimens 
examined through the kindness of the Berkeleys). The record of A. 
prismaticus from Oregon by Hartman and Reish (1950) is doubted 
since it is based on male stolons only. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms., stony-mud; off Point Barrow base, 5-70 fms., 
up to 7 miles from shore, on bottoms of stones, and various combina- 
tions of mud, gravel, stones (7 stations, 11 specimens). East Coast 
North America: Lagoon Pond, Martha's Vineyard, Massachusetts, 
pile scrapings, Pettibone, 1951. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Barents Sea, Novaya 
Zemlya. Also Iceland; Labrador to Massachusetts; Bering Sea to 
British Columbia ( ? Oregon) ; north Japan Sea. In low water to 267 
fathoms; sexual forms at surface. 

Genus Syllis Savigny, 1818 

Both species represented have the body elongate, slender, with 
numerous segments, flattened ventrally, arched dorsally, tapering 
slightly anteriorly and gradually posteriorly. Prostomium oval, wider 
than long; two palps wider basally, well separated except where their 
inner basal sides are fused (fig. 28, c). Dorsal cirri distinctly monili- 
form throughout body, alternately slightly longer and shorter. Ven- 
tral cirri digitiform, slightly shorter or longer than the parapodial 
lobes (fig. 28, d). Anterior end of extended proboscis with a smooth, 
chitinous rim, a single large, grayish dorsal tooth, with a ring of short, 
thick papillae, with a ring of 10 shorter papiUae more basally. Male 
and female stolons or epitokous forms (Chaetosyllis Malmgren, 1867) 
developed from the posterior segments of the body; when fully 
developed consisting of small bilobed (or tetralobed) bulbous "head" 
with two pairs of eyes, a dorsal pair and a larger ventral pair, with 
a pair of short moniliform antennae ; the first setigerous segment with 
neurosetae only, followed by a variable number of segments with 
additional long capillary swimming setae and usually a few unmodified 
posterior segments; stolons greatly distended with developing sex 
products; neurosetae, dorsal, ventral and anal cirri similar to those 
of stem form. 

Key to the species of Syllis from Point Barrow 

1. Compound neurosetae with short and long distal blades (fig. 28, /). Acicula 
of parapodia not particularly enlarged or protruding. Body segments very 
short S. cornuta 

Compound neurosetae with terminal blades differing very slightly in length 
(fig. 28, e) . From 1 to 4 very large acicula usually protrude somewhat from 
parapodial tip (fig. 28, d). Body segments longer S. fasciata 



MARESTE POLYCHAETE WORMS — PETTIBONE 253 

Syllis (Ehlersia) cornuta Rathke, 1843 

Figure 28, f 

Syllis (Ehlersia) cornuta Rathke, 1843, p. 164, pi. 7, fig. 12. — Malmgren, 1867, 

p. 43, pi. 7, fig. 45.— Southern, 1914, p. 37.— Fauvel, 1923, p. 267, fig. 100, 

g-i.— Augener, 1928, p. 718.— Monro, 1933, p. 34; 1937, p. 273; 1939b, p. 

387.— Wesenberg-Lund, 1947, p. 6; 1950a, p. 15; 1950b, p. 46; 1951, p. 36.— 

Zatsepin, 1948, p. 115, pi. 31, fig. 4. 
Chaetosyllis oerstedi Malmgren, 1867, p. 45, pi. 8, fig. 51 (epitokous form). 
Syllis oerstedi Th6el, 1879, p. 40, pi. 2, fig. 33; pi. 3, figs. 34, 35.— Annenkova, 

1938, p. 150.— Gorbunov, 1946, p. 38. 
Syllis quaternaria Moore, 1906a, p. 352, fig. (epitokous form). 
Syllis alternata Moore, 1908, p. 321, fig. (p. 324); 1909a, p. 321. — Annenkova, 

1938, p. 148.— Rioia, 1941, p. 691, pi. 3, figs. 1-9.— Berkeley and Berkeley, 

1948, p. 77, fig. 115. 
Syllis (Ehlersia) heterochaeta Moore, 1909a, p. 322, pi. 15, figs. 1-4. — Annenkova, 

1938, p. 148.— Rioja, 1941, p. 694, pi. 4, figs. 7-10; Berkeley and Berkeley, 

1948, p. 76, fig. 113. 
Ehlersia cornuta Hartman, 1945, p. 15. 
Typosyllis alternata Hartman, 1948, p. 21. 

Description. — Stem form: Length 9-45 mm., width without setae 
1-1.2 mm. Segments very short. Prostomium with two pairs of eyes, 
anterior pair larger, crescentric, with or without pair of small ocular 
spots anterior to lateral antennae. Moniliform dorsal cirri with about 
14-22 articles (11-40). Neurosetae all compound, with longer and 
shorter blades, the longer ones 2-4 times longer than the shorter ones, 
with jfine marginal fringe extending to near the tips (may give appear- 
ance of having a bidentate tip with a subterminal tooth) ; tips of short 
blades slightly hooked; tips of long, thin blades slightly hooked or 
with a slight knob (worn down?). In posterior segments, terminal 
blades of neurosetae not as long as in anterior segments. One speci- 
men with developing stolon of 41 segments beginning on segment 
75 of stem form. Color: In alcohol: (1) anterior segments dusky, 
then two narrow, reddish brown, transverse bands per segment on 
anterior and middle part of each segment; (2) rusty brown, with two 
darker bands per segment on part of body; (3) colorless. 

Male and female stolons (Chaetosyllis) : Length 8-19 mm., width 
without setae 1-1.2 mm., width including setae 4-5 mm., segments 
30-70 (1 unmodified anterior segment, 24-62 modified segments with 
swimming setae, 0-7 unmodified posterior segments); dorsal cirri 
alternately longer and shorter, with 11-18 articles (14-25 — Moore, 
1906); capillary setae much longer than the dorsal cirri; neurosetae as 
in posterior segments of stem form (distal blades not as long as in 
anterior region of stem form) ; some segments showed the characteristic 
banding of two narrow bands per segment. 



254 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Remarks. — The types of Syllis alternata Moore from Alaska and 
S. heterochaeta Moore from California were examined and are herein 
referred to S. cornuta (suggested previously by Monro (1933) for S. 
heterochaeta). The description of S. quaternaria Moore (1906), the 
epitokous form taken at the surface, Point Barrow, Alaska, agrees 
with that of the stolons of S. cornuta collected by Dr. MacGinitie in 
a plankton haul. 

New records. — Arctic Alaska: Stem form: Off Point Barrow base, 
18.3-123.5 fms., up to 15 miles from shore, on bottoms of stones, mass 
of worm tubes, and various combinations of mud, pebbles, stones, 
gravel, rocks, large perforated rocks, and worm tubes (17 stations, 97 
specimens). Sexual stolons: Off Point Barrow base, 1.6 miles from 
shore, vertical plankton haul of 13 fathoms tlirough hole in ice, March 
29, April 15, 1950 (12 specimens). West Coast North America: 
Strait of Juan de Fuca and Puget Sound, Washington, 30-107 fms., 
Pettibone. East Coast North America: Off Martha's Vineyard, 
86, 134, and 146 fms.; off Salem, Massachusetts, 35 fms., U. S. Fish 
Commission, 

Distribution. — Cosmopolitan. Widely distributed in the Arctic: 
Siberian and Alaskan Arctic, Greenland, Spitsbergen, Novaya Zemlya. 
Also Iceland, Norway to Madeira, Mediterranean; Red Sea, South 
Arabian coast, Persian Gulf; Indian Ocean (Zanzibar); off South 
Africa; Maine to North Carolina; north Japan Sea, Alaska to Panamd; 
South Pacific (Marquesas, Tahiti). In low water to 1,400 fathoms; 
sexual stolons in plankton. 

Syllis {Typosyllis) fasciata Malmgren, 1867 

FiauRE 28, o-e 

Syllis (Typosyllis) fasciata Malmgren, 1867, p. 43, pi. 7, fig. 47; pi. 8, fig. 52. — 
Augener, 1928, p. 719.— Fauvel, 1934b, p. 304.— Annenkova, 1934, p. 322; 
1938, p. 150.— Gorbunov, 1946, p. 38.— Wesenberg-Lund, 1947, p. 10, fig. 
2,a; 1950a, p. 16; 1950b, p. 47; 1951, p. 37.— Berkeley and Berkeley, 1948, 
p. 74, figs. 109, 110.— Zatsepin, 1948, p. 115, pi. 31, fig. 2. 

Syllis armillaris Treadwell, 1937, p. 28 (not Nereis armillaris Mtiller, 1776; in 
USNM). 

Description. — Stem form: Length 8-75 mm., width without setae 
0.7-3 mm. Many smaU ones present; many broken ones and a 
number with regenerating posterior ends. Segments not as short as 
in S. cornuta. Prostomium with two pairs of eyes, anterior pair 
larger, crescentric. Moniliform dorsal cirri with about 24 articles 
(20-40; 12-17 in young specimens). Parapodia each with one to 
four large, pointed acicula usually protruding beyond distal tips of 
setal lobes. Neurosetae all compound, with terminal blades rather 
short, hooked, with tips entire; some setae may have blades broken 



MARINE POLYCHAETE WORMS — PETTIBONE 255 

off (not to be confused with simple setae). Color: In alcohol: (1) 
pigmented with reddish-brown, one wide band per segment (darker 
on posterior part of band) ; more posteriorly two bands per segment, 
lighter anterior and darker posterior ones; then one band per segment; 
then without color (when mature, banded color pattern on the seg- 
ments of the developing sexual stolon); (2) colorless, particularly in 
small specimens; (3) uniform brown mottling on anterior third of 
body (specimens from Washington). In life: Reddish rusty brown 
bands, one wide band or two narrower bands per segment, especially 
on the anterior fourth and posterior fourth (latter a developing stolon, 
with wide, orange-red bands). One specimen with developing stolon 
beginning on segment 71. 

Sexual stolons (Chaetosyllis) : Two male stolons, 34-37 segments 
(one unmodified anterior segment, 31-33 modified segments with 
swimming setae, 2 or 3 small, unmodified posterior segments); long 
swimming notosetae subequal to dorsal cirri; dorsal cirri of first 
unmodified segment broken off at cirrophores; darkly pigmented 
reddish brown dorsaUy and ventrally. 

New Records. — Arctic Alaska: Stem form: Off Point Barrow base, 
18.3-123.5 fms., up to 15 miles from shore, on bottoms of mud, stones, 
worm tubes, from breaking rocks and bryozoans, from interstices 
between pebbles and gravel covering tunicate, Molgula sp., from 
various combinations of mud, pebbles, stones, gravel, rocks, large 
perforated rocks, worm tubes, and shells (21 stations, 206 specimens). 
Male stolons: Off Point Barrow base, 1.6 miles from shore, vertical 
plankton haul of 13 fms. through hole in ice (March 29, 1950, 2 speci- 
mens). West Coast North America: Washington Sound, 17-21 
fms., Pettibone. Canadian Arctic: Center Foxe Basin, 25-31 fms., 
Bartlett, 1927. 

Distribution. — Widely distributed in the Ai'ctic: Siberian, Alaskan, 
and Canadian Ai-ctic, Baffin Bay, Greenland, Jan Mayen, Spitsbergen, 
Franz Josef Land, Novaya Zemlya, Kara Sea. Also Iceland, Faroes; 
Bering Sea to southern California; north Japan Sea, China. In low 
water to 378 fathoms; sexual stolons in plankton. 

Genus Sphaerosyllis Claparfede, 1863 

Represented by a single species from Point Barrow. 

Sphaerosyllis erinaceus Clapar^de, 1863 

Figure 28, m 

Sphaerosyllis erinaceus Claparfede, 1863, p. 45, pi. 13, fig. 38 (fide Fauvel). — 
Southern, 1914, p. 20. — Fauvel, 1923, p. 302, fig. 115, q-r. — Annenkova, 
1934, p. 322; 1938, p. 153.— ? Rioja, 1943, p. 211, figs. 1-6.— Gorbunov, 1946, 
p. 38.— Zatsepin, 1948, p. 116, pi. 31, fig. 7. 



256 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Sphaerosyllis latipalpis Levinsen, 1882, p. 244. — Augener, 1928, p. 722, pi. 11, 
fig. 3; 1939, p. 140.— Wesenberg-Lund, 1947, p. 13, fig. 4; 1951, p. 38. 

Sphaerosyllis longicauda Webster and Benedict, 1887, p. 720, pi. 3, figs. 35-39. — 
Eliason, 1920, p. 11, fig. 3. 

Oophylax minuta Treadwell, 1937, p. 29, figs. 6, 7. 

Brania sp. Hartman, 1944a, pi. 24, figs. 1, 2. 

Description. — Length 2-4.5 mm., width without setae 0.3-0.5 mm., 
segments 22-37. Body linear, tapering slightly anteriorly and 
posteriorly, oval in cross section. Body, including parapodia, covered 
with small papillae and incrusted with fine, granular material. Pro- 
stomium subrectangular, wider than long, with six black eyes, four 
larger ones (each with a lens, arranged in transverse line or slightly 
concave arc near middle of prostomium) and two small anterior ones. 
Antennae subequal, bulbous basally, narrower distally. Paired 
antennae lateral to anterior eye spots; median antenna more posterior, 
nearly in line with the four larger eyes. Palps short, wide, rounded 
anteriorly; they may project more anteriorly, extending about the 
same length as the prostomium, with basal halves fused and distal 
halves separated by a narrow groove, or they may project more ven- 
trally, extending only about half the length of the prostomium, and 
fused except for a small anterior indentation; ventrally the palps 
are not fused although they may be closely approximated, extending 
back to the level of the four large eyes. Tentacular segment not 
distinctly set off from the prostomium, with a single pair of tentacular 
cirri which are similar to the antennae. Dorsal cirri short, slightly 
longer than parapodial lobes and shorter than the setae, inflated 
basally, tapering to narrow tips; they may be similar in shape through- 
out body or they may be only slightly inflated basally in middle and 
posterior regions. Dorsal cirri lacking on setiger 2. Ventral cirri 
digitiform, slightly shorter than parapodial lobes. Anal cutI larger 
than dorsal cirri; anal segment with several larger papillae. Para- 
podial tip usually with a larger papilla. Neurosetae consisting of a 
single long, simple, tapering, curved upper seta, the rest compound 
with distal blades long, subequal, entire, hooked, and finely pectinate. 
Colorless in alcohol. Female (21 fathoms, September 9, 1948) with 
large, rounded eggs (larger in diameter than parapodial length) fastened 
to dorsal surface rather irregularly between setigers 8-24, one to four 
per segment, attached between neuropodia and dorsal cirri and just 
medial to the dorsal cirri; another female (36 fathoms, October 6, 
1949) with large, oval-shaped developing larvae (developing setae 
visible) attached to the dorsal surface; swimming setae lacking (with 
swimming setae in male — Fauvel, 1923; swimming setae m both male 
and female beginning on setiger 8 — Webster and Benedict, 1887; 
eggs on ventral base of feet — Wesenberg-Lund, 1947). 



MARINE POLYCHAETE WORMS — PETTIBONE 257 

Remarks. — The type of Oophylax minuta Treadwell from Foxe 
Channel was examined and is herein referred to Sphaerosyllis erinaceus; 
although not mentioned in the original description, the specimen has a 
median antenna (seen with difficulty on the prepared slide); neuro- 
setae include a simple seta in addition to the compound ones, setiger 
2 lacks dorsal cirri. The type specimens of S. longicauda Webster and 
Benedict from Eastport, Maine, were examined also but they are 
unsatisfactory, being hardened, shrunken, and covered with crystals; 
the description is fairly complete and agrees with S. erinaceus as 
does the description by Augener of S. latipalpis Levinsen. The 
record by Kioja of S. erinaceus from Mexico is questioned chiefly 
because of the position of the eyes. 

New records. — Arctic Alaska: Off Point Barrow base, 16.7-75.5 
fms., up to 8 miles from shore, on bottoms of stones and various 
combinations of gravel, stones, rocks, large perforated rocks, with 
bryozoans and hydroids (8 stations, 37 specimens). East Coast 
North America: Off Labrador, 25 fms.. Blue Dolphin Expedition, 
1949. Vineyard Sound, Massachusetts, surface, U. S. Fish Commis- 
sion, August 3, 1881. 

Distribution. — Widely distributed in the Arctic: Siberian , Alaskan, 
and Canadian Arctic, West Greenland, Spitsbergen. Also Iceland, 
Denmark, Ireland, English Channel, Baltic; Labrador to Mas- 
sachusetts; Bering Sea; north Japan Sea; ? Mexico. In low water 
to 75.5 fathoms; sexual forms at surface. 

Genus Exogone Oersted, 1845 

Both species represented are very small, slender, linear-elongate, 
tapered slightly anteriorly and more so posteriorly, flattened ventrally, 
arched dorsally. Body colorless, translucent (in alcohol). Pro- 
stomium oval to subrectangular, arched somewhat anteriorly, much 
wider than long; four eyes, with lenses, in trapezoidal arrangement; 
palps fused dorsally into a rounded or triangular mass, with or without 
a slight anterior notch; palps separated ventrally by wide depression. 
Tentacular segment short, more or less distinct, the anterior part of 
which may form a fold covering the posterior part of the prostomium. 
Dorsal cirri shorter than parapodial lobes, ovoid to somewhat conical. 
Ventral cirri digitiform, shorter than parapodial lobes. Neurosetae of 
three kinds: (1) upper one simple, slightly curved; (2) composite, 
with longer, slender, pointed blades; (3) composite, with short, hooked 
blades. Everted pharynx crowned with circle of papillae and single 
conical tooth. With swimming setae at maturity. Large eggs, 
embryos or young (up to five setigers, resembling the adult) attached 
to ventral surface of female. 



258 PROCEEDINGS OF THE NATIONAL MUSEUM vol, 103 

Key to the species of Exogone from Point Barrow 

I. Antennae subequal. Tentacular cirri rudimentary, reduced to small knobs 

(fig. 28, 0- Dorsal cirri lacking on setiger 2. Upper compound setae with 

rather short pointed blades E. naidina 

Median antenna more than 4 times length of lateral antennae (fig. 28, k). 
Tentacular cirri only slightly smaller than the lateral antennae or dorsal 
cirri. Dorsal cirri present on all setigerous segments. Upper compound 
setae with long pointed blades E. dispar 

Exogone naidina Oersted, 1845 

Figure 28, I 

Exogone naidina Oersted, 1845, p. 20, pi. 2. — Augener, 1939, p. 140. — Zatsepin, 
1948, p. 116, pi. 31, fig. 8. 

Exogone gemmifera Pagenstecher, 1862, p. 267, pi. 25; pi. 26, figs. 1, 2, 6-8. — 
Mcintosh, 1908, p. 151, pi. 59, figs. 5, 6.— Southern, 1914, p. 17.— Eliason, 
1920, p. 11.— Fauvel, 1923, p. 305, fig. 117, a-d.— Annenkova, 1934, p. 322; 
1938, p. 154.— Rioja, 1943, p. 223, figs. 38-46.— Gorbunov, 1946, p. 38.— 
Thorson, 1946, p. 35, fig. 8.— Berkeley and Berkeley, 1948, p. 79, fig. 118.— 
Hartman and Reish, 1950, p. 13. 

Description. — Length 3.5-5 mm., width without setae 0.3 mm., 
segments 32-36. Palps form rounded mass, 1.4-2 times length of 
prostomium. Antennae shorter than palps, subequal or median one 
slightly longer, clavate, inserted anterior to eyes in almost a straight 
line. Tentacular cirri very small, reduced to small knobs. Dorsal 
cirri lacking on setiger 2. Neurosetae of each parapodium consisting 
of one simple, cm-ved upper seta, one upper composite seta with 
longer pointed blade, and four or five composite setae with short, 
hooked blades. Female (21 fathoms, September 15, 1948) without 
special swimming setae, with large eggs attached to ventral surface 
(between segments 1 1-25, one to three per segment) medial to ventral 
cirri and between parapodia. Another female (57 fathoms, October 

II, 1949) with eggs on ventral surface of segments 13-21 (mostly 
four per segment) ; eggs large, nearly touching. (Swimming setae on 
females with internal ova, absent in females with ova attached to 
ventral surface, two eggs per segment — Thorson, 1946.) 

New records. — Arctic Alaska: Off Point Barrow base, 16.7-123.5 
fms., up to 16 miles from shore, on bottoms of rocks, stones, worm 
tubes, and various combinations of mud, gravel, stones, rocks, large 
perforated rocks, with worm tubes, bryozoans, hydroids; on female 
Hyas coarctatus alutaceus (14 stations, 82 specimens). 

Distribution. — Alaskan and Siberian Ai-ctic; Denmark, Ireland, 
Great Britain to Madeira, Mediterranean; Bering Sea to Mexico; 
north Japan Sea. In low water to 123.5 fathoms; sexual stages at 
surface. 



MARINE POLYCHAETE WORMS — PETTIBONE 259 

Exogone dispar (Webster, 1879) 

Figure 28, k 

Paedophylax dispar Webster, 1879a, p. 223, pi. 4, fig. 49; pi. 5, figs. 50-55; 1879b, 

p. 110; 1886, p. 138.— Sumner, 1913, p. 615. 
Paedophylax longiceps Verrill, 1879, p. 170; 1881, p. 320, pi. 12, fig. 2; 1882, p. 

370.— Andrews, 1891, p. 281. 
Paedophylax longicirris Webster and Benedict, 1887, p. 722, pi. 3, figs. 46-50. 
Exogone lourei Berkeley and Berkeley, 1938, p. 44, figs. 6-12; 1948, p. 79, fig. 117.— 

Rioja, 1941, p. 703, pi. 3, figs. 14-21; 1943, p. 224. 
Exogone dispar Hartman, 1942a, p. 11; 1944a, p. 338, pi. 24, fig. 9; pi. 25, fig. 5; 

1945, p. 16, pi. 2, figs. 7, 9, 10; 1951, p. 40. 

Description. — Anterior ends of 3 specimens only (up to 8 mm. long, 
45 setigers — Berkeley and Berkeley, 1938). Palps form a conical 
mass, about twice length of prostomium. Antennae inserted between 
and slightly in front of eyes; median antenna longer, usually reaching 
nearly to end of palps (may be not much longer than prostomium), 
lateral antennae short, ovoid. Tentacular cirri slightly smaller than 
lateral antennae and dorsal cirri. Dorsal cirri present on all setigerous 
segments, including setiger 2, Neurosetae on each parapodium con- 
sisting of one simple curved upper seta (may have a delicate thread 
coming off subterminally in some specimens), one to four composite 
setae with long terminal capillary blades, and three to nine composite 
setae with short, hooked blades. 

Remarks. — The specimens were compared with the cotypes of 
Paedophylax dispar from New Jersey. The types of P. longicirris 
Webster and Benedict from Eastport, Maine, were examined also 
and the peculiar hairlike process on some of the simple setae was 
noted. This also occurs on some of the simple setae of the specimens 
from Point Barrow. P. longiceps Verrill has previously been referred 
to P. dispar by Sumner (1913). The description of E. lourei Berkeley 
and Berkeley from British Columbia agrees with that of E. dispar. 

New records. — Arctic Alaska: Off Point Barrow base, 36-70 fms., 
up to 7.5 miles from shore, on bottoms of stones, gravel, large per- 
forated rocks (2 stations, 3 specimens) . West Coast North America: 
Strait of Juan de Fuca, Washington, shore, Pettibone, 1940. East 
Coast North America: Woods Hole region, Massachusetts, Petti- 
bone, 1950, 1951. Maine, Massachusetts, New Jersey, low water to 
14 fms., U. S, Fish Commission. 

Distribution. — Arctic Alaska to Mexico; Maine to southwestern 
Florida. In low water to 70 fathoms; sexual stages at surface. 

Genus Eusyllis Malmgren, 1867 

Both species represented are flattened ventrally, arched dorsally, 
tapering both anteriorly and posteriorly, very fragile, breaking up 



260 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

easily and losing their antennae and dorsal cirri readily. Prostomium 
subrectangular to suboval, wider than long; four eyes fairly large, in 
trapezoidal arrangement, the anterior pair larger; palps broad, thick, 
oval to squarish, fused at their bases only and well separated distally, 
and may be curled ventrally or longitudinally like flopping ears; 
lateral antennae about 1.5 times the prostomial length; median 
antenna about twice the length of the lateral antennae (fig. 28, g). 
Upper tentacular cirri as long as median antenna, ventral pair much 
shorter. Ventral cirri thick, oval, about same size and length as 
parapodial lobes or may be slightly shorter (first pair not enlarged, 
similar to the following). Neuropodia short and plump, with neuro- 
setae all composite except on the last setigers where there may be a 
simple, entire, upper seta and a simple, bidentate, lower seta; blades of 
composite setae rather short, hooked, distinctly bidentate, finely 
spinous on the cutting edge; shaft distally spinous (fig. 28, i). At 
maturity, with long capillary swimming setae. Proboscis with two 
rows of soft papillae, a proximal and a distal row of 10 papillae per 
row; inside the distal row, a chitinous lining provided with a large, 
triangular, dorsal tooth and a denticled, circular margin (fig. 28, g, h). 

Key to the species of Eusyllis from Point Barrow 

1. Cirrophores of dorsal cirri not prominent. Tentacular segment with a wide 
crescent-shaped nuchal lobe covering the posterior part of the prostomium 

(fig. 28, g) E. blotnstrandi 

Cirrophores of dorsal cirri prominent, as long as the neuropodia. Tentacular 
segment short, without nuchal lobe E. magnifica 

Eusyllis blomstrandi Malmgren, 1867 

Figure 28, g-i 

Eusyllis blomstrandi Malmgren, 1867, p. 40, pi. 6, fig. 43. — Fauvel, 1923, p. 293, 
fig. 112,h-m.— Augener, 1928, p. 721.— Annenkova, 1938, p. 153, fig. 6.— 
Wesenberg-Lund, 1947, p. 11, fig. 3; 1950a, p. 16; 1950b, p. 48; 1951, p. 37.— 
Berkeley and Berkeley, 1948, p. 84, fig. 126. 

Eusyllis monilicornis Malmgren, 1867, p. 41, pi. 6, fig. 44. — Verrill, 1881, p. 319. — 
Zatsepin, 1948, p. 116, pi. 31, fig. 6,b. 

? Eusyllis phosphorea Hartman, 1942a, p. 7; 1944a, pp. 334, 338, pi. 25, fig. 4. 

Syllis monilicornis Th6el, 1879, p. 41, pi. 3, fig. 39. 

Eusyllis tubifex Mcintosh, 1908, p. 173, figs.— Southern, 1914, p. 32 (not Syllis 
tubifex Gosse, 1855). 

Eusyllis bloemstrandi Friedrich, 1939, p. 122. 

Typosyllis collaris Hartman, 1948, p. 23, fig. 6,a-c. 

Description. — Length 7-32 mm., width without setae 0.8-1.2 mm., 
segments 50-124. Antennae, tentacular cirri, anterior dorsal cirri, 
and long anal cirri irregularly annulated, especially distally, more or 
less smooth basally; the more posterior dorsal cirri only indistinctly 
annulated or smooth. First few dorsal cirri may be longer than body 
width, especially the first pair which may be the longest appendage; 



MARINE POLYCHAETE WORMS — PETTIBONE 261 

rest of dorsal cii-ri shorter thaii body width, tapering gradually. 
Tentacular segment with wide, short, crescent-shaped collar or nuchal 
fold covering posterior part of prostomium, may be inconspicuous 
and protruding or rather low, somewhat contracted. Composite 
setae similar throughout, with distal blades short, difTering only 
slightly in length. Chitinous lining of proboscis light or dark gray 
or light amber-colored; dorsal tooth dark bluish gray; circular margin 
finely denticled. Epitokous female (Elson Lagoon, August 6, 1948) 
with large eggs massed in most of body segments, consisting of 16 
anterior unmodified setigers, 25 setigers with long swimming setae, 
and 5 unmodified setigers (swimming setae begin on setiger 13 — 
Wesenberg-Lund, 1947). Color: In alcohol: Colorless or tannish, 
slightly greenish anteriorly. In life: Orange or yellowish. May form 
definite hardened mucous tubes. Luminescent. 

Remarks. — The type of Typosyllis coUaris Hartman (1948) was 
examined and is herein referred to E. hlomstrandi; the dorsal cirri 
are not distinctly annulated throughout the body as they would be 
in a Typosyllis. There is nothing in Verrill's description of E. phos- 
phorea, dredged from St. George's Bank, to distinguish it from E. 
hlomstrandi; according to Hartman (1942, p. 7) the type is unsatis- 
factory. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; Point Barrow base, washed ashore; off Point 
Barrow base, 13.3-123.5 fms., up to 15 miles from shore, on bottoms 
of stones, masses of worm tubes, and various combinations of mud, 
pebbles, gravel, stones, rocks, large perforated rocks, with worm tubes, 
bryozoans, hydroids, shells, and on female crab Hyas coarctatus alu- 
taceus (24 stations, 175 specimens). Bering Sea: Eobert White, 
1879. West Coast North America : Washmgton and Puget Sounds, 
20-46 fms., Pettibone. East Coast North America: Off Labrador, 
15-30 fms.. Blue Dolphin Expeditions, 1949, 1951. 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, West Greenland, Spitsbergen, Franz Josef Land, 
Barents Sea, Novaya Zemlya, Kara Sea. Also Iceland, Faroes, Ire- 
land to Madeira, Mediterranean; Labrador to Maine; Bering Sea to 
Wasliington; north Japan Sea. In low water to 444 fathoms; epitokes 
at surface. 

Eusyllis magnifica (Moore, 1906) 

Pionosyllis magnifica Moore, 1906b, p. 223, pi. 10, figs. 9-11; 1908, p. 325.— 
Annenkova, 1938, p. 152. 

Description.— Atokous specimen 52 mm. long, 2.5 mm. in greatest 
width without setae, 152 segments (in pieces, complete?). Epitokous 
specimen (18 fathoms, September 16, 1948) broken up and not quite 
complete, with larger eyes, consisting of 161 segments, 99 mm. long. 



262 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i03 

4 mm. wide without setae, 9 mm. wide including long swimming setae. 
Two atokous specimens from Washington 43-48 mm. long, 1.5-2 mm. 
wide without setae. Median antenna, tentacular cirri and some of 
anterior dorsal cirri somewhat annulated, especially distaUy; rest of 
dorsal cirri smooth. Some of anterior dorsal cirri longer than body 
width; rest shorter than body width, subulate, thick basally, tapering 
distally, on prominent cirrophores which may be as long as parapodial 
lobes. Tentacular segment very short, without nuchal collar. Blades 
of neurosetae all rather short. In anterior parapodia, blades of upper 
neurosetae somewhat longer than the lower ones; on rest of parapodia, 
blades of neurosetae only slightly different in length. Chitinous lining 
of proboscis and dorsal tooth light to darker amber-colored or trans- 
parent; circular anterior border with coarse, triangular denticles (when 
transparent, difficult to see denticled border). Swimming setae begin 
on setiger 24 (on setiger 19 in specimen from Portage Bay, Alaska). 
Color : In alcohol : Flesh. 

Remarks. — The specimens were compared with the type of Piono- 
syllis magnifica Moore (1906) from Port Townsend, Washington. 
It is herein referred to Eusyllis since the composite neurosetae have 
rather short blades which differ only slightly in length, the proboscis 
has a proximal and a distal row of papillae, and the anterior circular 
border is denticled (difficult to see when the chitinous lining is 
transparent) . 

New records. — Arctic Alaska: Off Point Barrow base, 18.3 fms., 
pebbles-mud (1 station, 2 specimens). West Coast North America: 
Albatross surface station. Portage Bay, Alaska; Washington Sound, 
Pettibone. 

Distribviion. — Arctic Alaska to Washington; north Japan Sea. In 
16 to 302.8 fathoms; epitokes at surface. 

Genus Pionosyllis Malmgren, 1867 
Pionosyllis compacta Malmgren, 1867 

Figure 28, j 

Pionosyllis compacta Malmgren, 1867, p. 40, fig. 48. — Augener, 1928, p. 722. — 
Annenkova, 1934, p. 322. 

Description. — Anterior ends only of two small specimens, up to 
1.5 mm. wide without setae (8-10 mm. long, 1 mm. wide — Malmgren, 
1867). Body flattened ventraUy, arched dorsally, fragile, with seg- 
ments crowded. Prostomium subrectangular, wider than long; four 
eyes rather large, in trapezoidal arrangement; palps squarish, fused 
basally, widely separated distally. Tentacular segment without 
nuchal collar. Antennae, tentacular, and dorsal cirri smooth, without 
articulations, filiform to rather thick, breaking off readily; most of 



MARINE POLYCHAETE WORMS — PETTIBONE 263 

dorsal cirri shorter than the body width. Ventral cirri thick, pointed, 
almost as long as parapodial lobes. Neurosetae all composite, distal 
blades distinctly bidentate, finely spinous along cutting edge; blades 
of upper setae longer, becoming gradually shorter in lower part of 
bundle with upper ones about twice the length of lower ones. Proboscis 
with a distal row of 10 papillae; transparent chitinous lining with a 
translucent bluish-gray dorsal tooth, with distal rim smooth. Color: 
In alcohol: Irregularly dotted with black pigment both dorsally and 
ventrally. (Epitokous forms have capillary setae beginning on seg- 
ments 12-15 — Malmgren, 1867.) 

New records. — Arctic Alaska: Off Point Barrow, 36-78 fms., up 
to 15 miles from shore, on bottoms of rocks, stones, and worm tubes 
(2 stations, 2 specimens). 

Distribution. — Arctic Alaska; Spitsbergen; Bering Sea. In low 
water to 78 fathoms. 

Family Nereidae 

Prostomium distinct, with four eyes, two frontal antennae, two bi- 
articulated palps (fig. 30, a). Four pairs tentacular cirri; buccal or 
tentacular segment usually apodous and achaetous. Parapodia 
usually biramous (except first two pairs), with dorsal and ventral 
cylindrical cirri, two or three notopodial ligules and one neuropodial 
ligule (fig. 30, 6, c). Setae usually compound spinigers and falcigers 
(fig. 30, d-g). Pair of anal cirri. Proboscis eversible, with pair of 
terminal, horny, falcate jaws and usually with horny denticles or 
paragnaths arranged in eight groups on two rings, a basal or oral 
ring and a distal or maxillary ring (fig. 30, h, i). At maturity usually 
with a pelagic epitokous or heteronereis stage. 

Represented by a single genus and two species. 

Genus Nereis Cuvier, 1817 

Subgenus Nereis Linne, 1758 

Both species have the body smooth, elongate, cylindrical, at- 
tenuated posteriorly, with numerous segments. Prostomium of the 
typical nereid form. Tentacular segment achaetous, over twice the 
length of the following segments; tentacular cirri short, fairly uniform 
in length, the upper one only slightly longer than the others (may 
reach setiger 2). Parapodia biramous, with two subequal noto- 
podial ligules (remaining so throughout the body, the upper one not 
getting larger or longer than the lower one), a single neuropodial 
ligule ventral to the conical setigerous neuropodial lobe (fig. 30, h, c). 
Notosetae homogomph spinigers (fig. 30, g); in more posterior seg- 
ments with some homogomph falcigers with blunt asymmetrical 
reduced appendages (fig. 30, e). Neurosetae homogomph and 



264 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

heterogomph spinigers (fig. 30, /) and heterogomph falcigers with 
hooked end-pieces (fig. 30, d). Proboscis with paragnaths all 
chitinous, conical, separated in groups, occurring on both rings 
(fig. 30, h, i). Distal ring of paragnaths: Area II (dorsolateral) — 
oblique group of two or three rows; area III (medio ventral) — small 
transverse group; area IV (ventrolateral)— arched mass. Basal ring 
of paragnaths: Area V (mediodorsal) — 0. At maturity, with epito- 
kous stage, eyes enlarged; 14-19 anterior prenatatory setigers; 
modified natatory setigers with parapodial lobes compressed, flat- 
tened anteroposteriorly, developing foliaceous lamellar plates, with 
numerous paddlelike swimming setae; in males, first seven dorsal 
cirri enlarged and club-shaped, first five or six ventral cirri modified, 
and dorsal cirri of natatory setigers crenulate on lower margin; in 
females, dorsal and ventral cutI not modified or crenulate. 

Key to the species of Nereis (Nereis) from Point Barrow 

1. Parapodial ligules short, thick, evenly rounded (fig. 30, b). Body uniformly 
purplish or reddish brown, not banded. Paragnaths of distal ring: Area 
I (mediodorsal) — 2 in tandem (rarely 1 or 3); paragnaths of basal ring: 
Area VI (dorsolateral) — 4 in square or cross (rarely 3 or 5); areas VII- 
VIII (ventral) — 1-2 irregular rows of larger paragnaths, vnth. several rows 
of small ones, diminishing in size posteriorly (2-3 irregular rows of subequal 
paragnaths in subspecies occidentalis). Male epitokes with 16 and females 

with 17-19 prenatatory setigers N. (N.) pelagica 

Parapodial ligules triangular to conical, gradually tapering to a broad tip 
(fig. 30, c). Body transversely banded reddish brown or violet. Parag- 
naths of distal ring: Area I — or 1; paragnaths of basal ring: Area VI — 
6-10 or more in oval mass; VII-VIII — continuous row of larger parag- 
naths followed by a wide band of small subequal ones (fig. 30, hy i). Mais 
epitokes with 14 and females with 16 prenatatory setigers N. (N.) zonata 

Nereis {Nereis) pelagica Linne 1758 

Figure 30, a, b 

Nereis pelagica Linn6, 1758, p. 654. — Malmgren, 1867, p. 47, pi. 5, fig. 35. — 
Webster and Benedict, 1884, p. 718; 1887, p. 724.— Moore, 1903, p. 431.— 
Mcintosh, 1908, p. 268, figs.— Sumnsr, 1913, p. 619.— Southern, 1914, p. 
80.— Eliason, 1920, p. 29.— Chamberlin, 1920, p. 14.— Fauvel, 1923, p. 336, 
fig. 130, a-f.— Augener, 1928, p. 712.— Monro, 1930, p. 106.— Annenkova, 
1934, p. 322; 1937, p. 162; 1938, p. 158.— Treadwell, 1937, p. 29.— Okuda, 
1939, p. 230.— Friedrich, 1939, p. 123.— Hartman, 1940, p. 225, pi. 35, fig. 
52; 1944a, p. 335; 1948, p. 26.— Berkeley and Berkeley, 1943, p. 130; 1948, 
p. 66, fig. 96, a-b.— Thorson, 1946, p. 64, fig. 29.— Zatsepin, 1948, p. 119, 
pi. 30, fig. 2.— Wesenberg-Lund, 1950a, p. 19, pi. 6, figs. 26a, 27a; 1950b, 
p. 53; 1951, p. 39. 

Heteronereis grandifoUa Rathke, 1843, p. 155, pi. 7, figs. 13, 14. — Malmgren, 
1865, p. 108, pi. 11, fig. 15 (part — female heteronereid, not male). 

Nereis neonigripes Hartman, 1936b, p. 471, fig. 48. 

Nereis arctica Treadwell, 1937, p. 30. 

Nereis pelagica var. occidentalis Hartman, 1945, p. 20, pi. 4, figs. 1-6; 1951, p. 46. 

Nereis pelagica neonigripes Hartman and Reish, 1950, p. 17. 



MARINE POLYCHAETE WORMS — PETTIBONE 265 

Description. — Length 60-155 mm., width 5-14 mm. See key. 
Color: In alcohol: Iridescent, uniformly purplish or reddish brown 
(narrow white intersegmental lines) ; ligules of parapodial lobes may 
be darkly pigmented (var. neonigripes) ; tips of dorsal cirri may be 
black. 

New records. — ^Arctic Alaska: Off Point Barrow base, up to 12.1 
miles from shore, 18-87 fms., on various combinations of stones, 
gravel, rocks, large perforated rocks (8 stations, 10 specimens). 
Spitsbergen: Spitsbergen Sea, U. S. S. Alliance, August, 1881, sm-face 
(heteronereid) . Bering Sea : Albatross Sta. at Nikolski, Bering Island, 
1892, and Sta. 3496, 56°32' N., 169°45' W., 41 fms., 1893; St. George 
Island, Pribilofs, 1913, 1914, low tide and 30 fms., and St. Paul Island, 
Pribilofs, lagoon reef, 1915, Hanna; St. Paul Island, Wm. Palmer, 
1890. Southwestern Alaska: Albatross Sta., Unalaska, Aleutians, 
1888; Akutan Pass, Chica Islands, 1872; Easka Harbor, sandy- 
mud, 9-12 fms., 1873; Anchorage, Big Koniuji Islands, Shu- 
magin, 6-20 fms., sand and rocks; Port Moller, Alaska Peninsula, 
beach to 17 fms., sand, 1874; KiUisnoo, 1897; Chiniak Bay, Kodiak 
Island, 1880; all collected by Dall. Gulf of Alaska: Albatross Sta., 
at Observation Island, Cordova, Alaska, 1914. Southeastern 
Alaska: Sitka, 1932. West Coast North America: Albatross 
Sta. 89a, Denman, British Columbia, 1914, and Sta. 3466, off Wash- 
ington, 48°18'30" N., 123°22' W., 56 fms., 1891. Washington and 
Puget Sounds, low tide to 166 fms, (very common), Pettibone. East 
Coast North America: Off Labrador, 7-60 fms.. Blue Dolphin 
Expeditions, 1949, 1951; Woods Hole region, intertidal and dredged, 
Pettibone, 1950, 1951; Gulf of St. Lawrence, New Brunswick, Nova 
Scotia, Bay of Fundy, Maine, Massachusetts, Ehode Island, Con- 
necticut, Long Island Sound, shore to 250 fathoms and surface, U. S. 
Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenlaad, Spitsbergen, Barents 
Sea, Novaya Zemlya, Kara Sea. Also Iceland, Faroes, Norway to 
Mediterranean; Hudson Bay to Long Island Sound; Bering Sea to 
Panamd; north Japan Sea, Japan; South Atlantic (Tristan da Cunha, 
Kerguelen, Magellan Straits). In low water to 609 fathoms; epitokes 
at surface. Variety occidentalis Hartman: North Carolina, Florida, 
Louisiana, Texas, Puerto Rico; intertidal. 

Nereis (Nereis) zonata Malmgren, 1867 

Figure 30, c, h, i 

Heteronereis grandifolia Malmgren, 1865, p. 108, pi. 11, fig. 16 (part — male 
heteronereid, not female; not H. grandifolia Rathke, 1843). 

Nereis zonata Malmgren, 1867, p. 46, pi. 5, fig. 34.— Th6el, 1879, p. 42.— Wir^n, 
1883, p. 402.— Ditlevsen, 1911, p. 419, pi. 28, fig. 6; pi. 30, figs. 18, 22.— 



266 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Fauvel, 1914, p. 177, pi. 14, figs. 1-17; 1923, p. 338, fig. 130, g-n (part).— 
Augener, 1928, p. 713.— Annenkova, 1931, p. 203; 1932, p. 134; 1934, p. 
322; 1937, p. 162; 1938, p. 158.— Friedrich, 1939, p. 123.— Berkeley and 
Berkeley, 1943, p. 130.— Gorbunov, 1946, p. 38.— Zatsepin, 1948, p. 119, 
pi. 30, fig. 3. — Hartman, 1948, p. 25. — Hartman and Reish, 1950, p. 17. — 
Wesenberg-Lund, 1950a, p. 20, pi. 6, figs. 26b, 27b; 1950b, p. 55; 1951, p. 41. 

Description. — Length 30-125 mm., width 7 mm. See key. Color: 
In alcohol: Reddish or rusty brown or violet wide transvei-se bands; 
anterior fourth of segment Avithout color. 

Remarks. — Nereis procera Ehlers, 1868, originally described from 
the Gulf of Georgia, British Columbia, has been referred to N. zonata 
by Fauvel, 1914. In A^. procera the hodj is long, slender, extremely 
attenuated posteriorly; the notopodial ligules of the anterior setigers 
are triangular, subequal; those of the median and posterior regions 
have the upper notopodial ligules larger than the lower one (may be 
nearly twice the size) ; the paragnaths of areas VII and VIII are in a 
wide, continuous area with no especially enlarged anterior row; male 
epitokes with 51 and females with 59 prenatatory setigers. It may be 
that the variety procera of Fauvel refers to a different species from that 
of N. procera Ehlers. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagooa near 
Point Barrow, 13 fms., gravel; off Point Barrow base, 21-78 fms., on 
bottoms of gravel, stones, small rocks, with shells and worm tubes (8 
stations, 12 specimens). Canadian Arctic: Ellesmere Island, U.S.S. 
Alert, 1950, J. Peter Johnson (epitoke). East Greenland: 74°04' N., 
17°58' W., 120 fms., Norcross-Bartlett Expedition, 1931. East 
Coast North America: Off Labrador, 9-100 fms,, mud. Blue Dolphin 
Expeditions, 1950, 1951. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Baffin Bay, Davis Strait, Greenland, Norway, 
Spitsbergen, Franz Josef Land, Barents Sea, Novaya Zemlya, Kara 
Sea. Also Iceland, Faroes; Hudson Bay to Labrador; Bering Sea to 
Oregon; north Japan Sea. In low water to 439 fathoms; epitokes at 
surface. Variety ? procera Fauvel (1914, 1923; not Ehlers, 1868): 
English Channel to Mediterranean. Variety persica Fauvel: Red 
Sea, South Arabian Coast, Persian Gulf, Indian Ocean. 

Family Nephtyidae 

Body elongated, subtetragonal in cross section. Prostomium small, 
somewhat flattened, angulate, with four small antennae (fig. 30, j). 
First segment rudimentary, with setae and usually with short dorsal 
and ventral tentacular chri. Parapodia bu-amous, with rami well 
separated, with smaU dorsal chri on lower sides of notopodia; with 
branchiae (interramal chri) between the two rami, cirriform, sickle- 
shaped or foliaceous; with simple setae arranged in fan-shaped bundles; 



MARINE POLYCHAETE WORMS — PETTIBONE 267 

ventral cirri short, conical, or foliaceous; with anterior and posterior 
lamellae more or less developed (fig. 30, k-^). A single anal cirrus. 
Proboscis eversible, strong, muscular, a pah- of horny jaws wdthin the 
pharynx, with rows of soft papillae and a crown of terminal, bifid 
papillae. Move rapidly with undulating motion; usually in sand 
or mud. 

Represented by a single genus and four species. 

(ienus Nephtys Cuvier, 1817 

The four species represented have numerous segments (65-150), 
tapered posteriorly and slightly anteriorly. Prostomium pentagonal 
to quadrate, with antennae conical, subequal, or more ventral pair 
slightly larger, without eyes, with pair of nuchal papillae at postectal 
margins (may be inverted or everted; fig. 30, j). Tentacular or first 
segment with both dorsal and ventral tentacular cu-ri. Parapodial 
rami with anterior lamellae rudimentary or lacking, with posterior 
lamellae developed m varying degrees, with branchiae recurved (con- 
vex side toward lateral side of body, fig. 30, Ar-n). Ventral cu-ri short, 
conical. Proboscis with 22 long terminal papillae (10 pairs bifid, and 
a single dorsal and ventral one), with 22 longitudinal rows of sub- 
terminal papillae, four to eight per row, decreasing in size basally. 

Key to the species of Nephtys from Point Barrow 

1. Branchiae cirriform, sickle-shaped, with dorsal 'curi digitiform (fig. 30, I, n). 

Proboscis with a long median dorsal papilla 2 

Branchiae foliaceous or cirriform and inflated basally, rudimentary or absent 
on last 15-30 segments, with dorsal cirri flattened, triangular, pointed (fig. 
30, k, m). Proboscis without a long median dorsal papilla; proximal surface 
of proboscis ridged (may be low tubercles in N. discors) 3 

2. Branchiae begin on setigers 3 or 4, continuing to almost posterior end; acicular 

lobes rounded or sUghtly bilobed, especially anteriorly (fig. 30, I). Neuro- 
podial postsetal lamellae extend well beyond the acicular lobes, bilobed or 
irregularly sinuous. Setae very long, flowing. Proximal surface of pro- 
boscis smooth or furrowed N. longosetosa 

Branchiae begin on setigers 5-8, rudimentary on last 20-30 segments. Acicular 
lobes deeply bilobed in anterior and median regions (fig. 30, n) . Neuropodial 
postsetal lamellae about same length as or only slightly surpassing the 
acicular lobes. Setae shorter. Proximal surface of proboscis tubercled. 

N. ciliata 

3. Branchiae begin on setiger 6, inflated basally, digitiform, recurved distally 

(fig. 30, m). Acicular lobes unequally bilobed, rounded in posterior region. 
Postsetal lamellae of anterior and median regions large, foliaceous, extending 
well beyond the acicular lobes; in posterior region, extend only slightly 

beyond N. discors 

Branchiae begin on setigers 8-14, thick, wide, foliaceous (fig. 30, k). Acicular 
lobes rounded to conical. Postsetal lamellae shorter than, as long as, or 
only slightly longer than the acicular lobes N. paradoxa 

261112—54 5 



268 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Nephtys longosetosa Oersted, 1843 

FlGUBE 30, I 

Nephtys longosetosa Oersted, 1843, p. 195, pi. 6, figs. 75, 76. — Hartman, 1944a, 
p. 339, pi. 15, fig. 7. 

Nephthys longisetosa not Malmgren, 1865, p. 106, pi. 12, fig. 20. — Eliason, 1920, 
p. 32. 

Nephthys coeca Murdoch, 1885, p. 153 (part; includes var. longisetosa; not Nereis 
caeca Fabricius, 1780). 

Nephthys longosetosa Fauvel, 1923, p. 367, fig. 143, f-h. — Monro, 1928, p. 82. — 
Annenkova, 1937, p. 163; 1938, p. 162.— Friedrich, 1939, p. 123.— Berkeley 
and Berkeley, 1943, p. 130; 1948, p. 52, fig. 76.— Gorbunov, 1946, p. 38.— 
Wesenberg-Lund, 1950a, p. 21; 1950b, p. 60; 1951, p. 44. 

Nephthys ciliata forma longosetosa Augener, 1928, p. 698. 

Nephthys ciliata Treadwell, 1937, p. 28 (part; not Nereis ciliata Muller, 1789). 

Description. — Length up to 170 mm., width without setae 6 mm. 
Tentacular cirri subequal to antennae. Notopodial postsetal lamellae 
unequally bilobed (a rather large upper lobe and small lobe just above 
the dorsal cirri), extending slightly or not beyond the acicular lobes. 
Neuropodial postsetal lamellae of anterior region rounded, entire, 
extending beyond the acicular lobes; those of the median and posterior 
regions large, bilobed or irregularly sinuous on ventral side, extending 
well beyond the acicular lobes. Color: In alcohol: Without color, 
or dusky, especially anteriorly. 

New records. — Arctic Alaska: Eluitkak, Elson Lagoon near Point 
Barrow; Point Barrow base, washed ashore; off Point Barrow base, 
1.7-25 fms., on bottoms of gravel, stones, and mud (7 stations, 16 
specimens); Cape Lisburne, 5-7 fms., Dall. Bering Sea: 60°16' N., 
167°41' W., Stoney, 1884; 66°45' N., 166°35' W., 10 fms., Dall, 1880. 
Canadian Arctic: Duckett Cove, Hurd Channel, MelvUle Peninsula, 
Foxe Channel, Bartlett, 1933. East Coast North America: Off 
Labrador, 8 fms.. Blue Dolphin Expedition, 1949. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Franz Josef Land, 
Barents Sea, White Sea, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes to France; to Maine; to Lower California and Panamd; north 
Japan Sea; Strait of Magellan. In low water to 528 fathoms. 



Figure 30. — Nereidae: a, Nereis pelagica, dorsal view anterior end; b, same, middle para- 
podium; c. Nereis zonata, middle parapodium; d, compound heterogomph falciger; e, 
compound homogomph falciger; /, compound heterogomph spiniger; g, compound homo- 
gomph spiniger; h. Nereis zonata, dorsal view extended proboscis showing areas of parag- 
naths; i, same, ventral view. Nephtyidae: /, Nephtys paradoxa, dorsal view anterior 
end; k, same, middle parapodium, anterior view; /, Nephtys longosetosa, middle para- 
podium, anterior view; m, Nephtys discors, middle parapodium, anterior view; n, Nephtys 
ciliata, middle parapodium, anterior view. (For explanation of symbols, see p. 210.) 



MARINE POLYCHAETE WORMS — PETTIBONE 



269 




FiGtrRE 30. — For explanation see facing page. 



270 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Nephtys ciliata (Miiller, 1789) 

Figure 30, n 

Nereis ciliata Miiller, 1789, p. 14, pi. 89, figs. 1-4. 

Nephthys ciliata Malmgren, 1865, p. 104, pi. 12, fig. 17. — Webster and Benedict, 

1887, p. 709.— Moore, 1903, p. 433.— Eliason, 1920, p. 31.— Fauvel, 1923, 

p. 371, fig. 145, a-b; 1933, p. 39.— Augener, 1928, p. 699.— Gustafson, 1936, 

p. 7.— Treadwell, 1937, p. 28 (part) .— Annenkova, 1937, p. 164; 1938, p. 

162.— Friedrich, 1939, p. 123.— Gorbunov, 1946, p. 38.— Thorson, 1946, p. 

69, figs. 32, 33.— Berkeley and Berkeley, 1948, p. 55, fig. 82.— Wesenberg- 

Lund, 1950a, p. 21; 1950b, p. 59; 1951, p. 45. 
Nephthys caeca Murdoch, 1885, p. 153 (part; includes var. ciliata). — Moore, 1911, 

p. 243 (not Nereis caeca Fabricius, 1780). 
Nepthys ciliata Chamberlin, 1920, p. 9. 
Nepthys hudsonica Chamberlin, 1920, p. 10, pi. 2, figs. 4-6. 
Nephtys ciliata Hartman, 1944a, pp. 335, 339, pi. 15, fig. 9 (as N. incisa, not fig. 

10 {=N. caeca)); 1950, p. 95. 

Description. — Length 100-300 mm., width without setae 9-11 mm. 
Tentacular cirri subequal to antennae. Acicular lobes deeply bilobed 
in anterior and median regions; rounded or slightly bilobed in posterior 
region. Postsetal lamellae of both notopodia and neuropodia rounded, 
entire, short, about same length as or only slightly surpassing the 
acicular lobes. Color: In alcohol: White, tannish, or anterior third 
reddish brown or buff. (According to G. E. MacGinitie, the pellets 
of this worm indicate that it is a detritus feeder.) 

New records. — Arctic Alaska: Off Point Barrow aero radio mast, 
10 fms.; off Point Barrow base, up to 8 miles from shore, 6-75.5 fms., 
on bottoms of mud and various combinations of mud, pebbles, gravel, 
stones, rocks, large perforated rocks, and shells (18 stations, 37 speci- 
mens). Siberia: Plover Bay, Dall, 1880. Alaska: Chiachi Island, 20 
fms., mud; Round Island, Coal Harbor, 6-8 fms., mud, 1872; Port 
Mulgrove, Yakutat Bay, 6-40 fms., 1874; Port Etches, 12-18 fms., 
1874; aU collected by DaU. West Coast North America: Wash- 
ington and Puget Sounds, low water to 83 fms. (common), Petti- 
bone. East Coast North America: Off Labrador, 6-95 fms.. Blue 
Dolphin Expeditions, 1949, 1950, 1951; off Nova Scotia, Bay of 
Fundy, Maine, Massachusetts, 14-110 fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
aud Canadian Arctic, Greenland, Spitsbergen, Norway, Barents Sea, 
White Sea, Novaya Zemlya, Kara Sea. Also Iceland, Faroes to 
France; to Massachusetts; to Southern California; to Japan, China. 
In low water to 500 fathoms. 

Nephtys discors Ehlers, 1868 

Figure 30, m 

Nephthys discors Ehlers, 1868, p. 626, pi. 23, figs. 39, 40.— Verrill, 1873, p. 103.— 

Webster and Benedict, 1887, p. 709. 
Nephtys discors Hartman, 1938c, p. 9, pi. 1, figs. 2, 3; 1950, p. 96. 
Nephtys rickettsi Hartman, 1938b, p. 153; 1950, p.',97. 



MARINE POLYCHAETE WORMS — PETTIBONE 271 

Description. — Length 115-300 mm., width without setae 9-12 mm. 
Dorsal pair tentacular cuTi about half as large as ventral pair. 
Branchiae begin on setiger 6, at first a small swelling on the lower side 
of the base of the flattened, triangular dorsal cirrus, gradually becom- 
ing larger; where best developed, they are inflated on the basal half, 
with the distal half more slender, digitiform and slightly recurved or 
hooked; they are smaller and rudimentary only on posterior half of 
body. Notopodial acicular lobes of anterior and middle region 
unequally bilobed, the upper one wider and shorter and the ventral 
one narrower and longer; rounded in posterior region. Neuropodial 
acicular lobes slightly bilobed in anterior region; very low and rounded 
in middle and posterior regions (not very distinct from posterior 
lamellae). Color: In alcohol: Brownish. 

Remarks. — The type of N. rickettsi Hartman from Alaska was 
compared with a specimen of A^. discors from Bay of Fundy and is 
herein referred to the latter species. 

New records. — Arctic Alaska: Point Barrow base, washed ashore; 
off Point Barrow base, up to 12.1 miles from shore, 24.7-123.5 fms., 
on bottoms of mud, gravel-mud, stones and large perforated rocks, 
and mass of worm tubes (5 stations, 5 specimens). West Coast 
North America: Puget and Washington Sounds, 70-83 fms., mud 
(3 specimens), Pettibone, 1938, 1939. East Coast North America: 
Bay of Fundy, U. S. Fish Commission. 

Distribution. — Alaskan Arctic to southern California; Bay of Fundy 
to Maine. In 24-268 fathoms. 

Nephtys paradoxa Malm, 1874 

Figure 30, j, k 

Nephthys paradoxa Malm, 1874, p. 78, pi. 1, fig. 2. — Fauvel, 1923, p. 375, fig. 
146, f-i.— Augener, 1928, p. 701.— Gustafson, 1936, p. 7.— Annenkova, 1938, 
p. 163.— Friedrich, 1939, p. 123.— Berkeley and Berkeley, 1943, p. 130.— 
Wesenberg-Lund, 1950a, p. 22; 1950b, p. 61; 1951, p. 47. 

Nephthys phyllobranchia Mcintosh, 1885, p. 164, pi. 26, fig. 10; pi. 27, fig. 3; pi. 
14, A, figs. 12, 13. 

? Nephthys brachycephala Moore, 1903, p. 431. — Annenkova, 1937, p. 164; 1938, 
p. 162.— Zatsepin, 1948, p. 121, pi. 30, fig. 9. 

Nephtys paradoxa Hartman, 1944a, pp. 335, 339, pi. 15, fig. 6; 1950, p. 111. 

Nephtys phyllobranchia Hartman, 1950, p. 111. 

Description. — Length 150-200 mm., width without setae 8-13 mm. 
Dorsal pair tentacular cirri reduced to a tubercle, smaller than ventral 
pair. Branchiae begin on setigers 11-13 (8-14), very short, triangular 
at first, gradually becoming larger; where best developed, they are 
flat, wide foliaceous, weakly to distinctly concave; they are small, 
nearly absent on the last 15-30 segments. The anterior acicular 
lobes are rounded; those of the middle and posterior regions are 
conical, pointed. The postsetal lamellae in the anterior region are 



272 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

slightly longer than the acicular lobes; they are shorter than the 
conical acicular lobes in the middle and posterior regions. Color: 
In alcohol : Dusky, especially anteriorly. 

Remarks. — Augener (1928) indicated that he was of the opinion 
that N. phyllobranchia Mcintosh should be referred to A^, paradoxa; 
Hartman (1950) stated that they are separable by only slight differ- 
ences. The type of A^. brachycephala Moore from Japan is unsatis- 
factory (dried, when received); the description is in agreement with 
N. paradoxa. 

New records. — Arctic Alaska: Off Point Barrow base, up to 12.1 
miles from shore, 23.5-123.5 fms., on bottoms of mud and mass of 
worm tubes (2 stations, 3 specimens). 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Norway, Barents Sea, 
Kara Sea. Also Iceland, Faroes to France; off Delaware; Japan. 
In 3.3-4,001 fathoms. 

Family Glyceridae 

Body rounded, elongated, attenuated at both ends, not divided 
into two regions, either only uniramous or only biramous. Segments 
bi- or tri-annulate. Prostomium sharply conical, wide basaUy, with 
a pair of nuchal organs, transversely annulated, ending in four minute 
antennae distally (fig. 31, a). Small, globular dorsal cirri; larger 
ventral cirri (fig. 31, b, c); two anal cirri. Branchiae present or 
absent. Notosetae simple capillaries; neurosetae compound spinigers. 
Proboscis eversible, long, clavate, with numerous, small papillae 
(proboscidial organs) and four dark, falcate, horny jaws each with an 
attached, embedded, rodlike aileron (fig. 31, d). Epitoky affects 
entke individual (parapodial lobes become more elongate, setae longer 
and more numerous). 

Represented by a single species from Point Barrow. 

Genus Glycera Savigny, 1818 
Glycera capitata Oersted, 1843 

FiGUBE 31, a-d 

Glycera capitata Oersted, 1843, p. 196, pi. 7, figs. 87, 88, 90-94, 96, 99.— Ehlers, 
1913, p. 603.— Fauvel, 1923, p. 385, fig. 151, a-e.— Augener, 1928, p. 734.— 
Monro, 1930, p. 115; 1936, p. 114.— Annenkova, 1937, p. 165; 1938, p. 164.— 
St0p-Bowitz, 1941, p. 186, fig. 1; 1948b, p. 4.— Hartman, 1944a, pp. 336, 
339; 1948, p. 28; 1950, p. 76, pi. 11, figs. 1-4.— Gorbunov, 1946, p. 38.— 
Zatsepin, 1948, p. 117, pi. 31, fig. 9.— Berkeley and Berkeley, 1948, p. 38, 
fig. 52. — Hartman and Reish, 1950, p. 20. — Wesenberg-Lund, 1950a, p. 23; 
1950b, p. 64; 1951, p. 49. 

Glycera setosa Oersted, 1843, p. 198, pi. 7, figs. 89, 95, 97 (epitokous form). 



MARINE POLYCHAETE WORMS — PETTIBONE 273 

Glycera nana Johnson, 1901, p. 411, pi. 10, fig. 103.— Berkeley and Berkeley, 

1948, p. 37, figs. 50, 51. 
Glycera lapidum Fauvel, 1914, p. 205; 1923, p. 386, fig. 151, /-m; 1934a, p. 41.— 

Eliason, 1920, p. 36, fig. 6.— Treadwell, 1937, p. 32.— St0p-Bowitz, 1941, 

p. 191, fig. 2; 1948b, p. 7. 

Description. — Length 80 mm., width without setae 6 mm. (length 
up to 150 mm. — Hartman, 1950). Segments usually triannulate 
(may be biannulate, a slightly longer anterior ring at the level of the 
parapodia and a shorter posterior ring; the former may be sub- 
divided, thus triannulate; the latter may also be somewhat divided, 
thus tetra-annulate) . Conical prostomium with 8-16 annuli (often 
difficult to count accurately). Dorsal cirri small, globular, inserted 
well above the parapodia. Ventral cirri wide, conical. Parapodia 
biramous throughout; notopodia represented by a short, conical 
presetal lobe and a small bundle of simple capillary setae; neuropodia 
with a longer, conical presetal lobe and a fan-shaped group of com- 
pound spinigerous setae; a single short, wide, rounded postsetal lobe 
common to both notopodia and neuropodia (thus two conical an- 
terior lobes and a single postsetal lobe). The relative length of the 
parapodial lobes may be variable, associated with the epitokous 
condition, as indicated by Hartman (1950). Proboscis with two 
kinds of papillae, numerous, long, cylindrical, slender ones, and 
fewer, larger, short, subspherical ones; aileron of jaws with a long 
lateral prolongation. Color: In alcohol: Brownish. 

Remarks. — Monro (1930) referred Qlycera lapidum Quatrefages to 
G. capitata; they have been separated on minor and apparently 
variable characters; Fauvel (1914, 1923) regarded them as mere 
varieties. Hartman (1950) referred G. nana Johnson to G. capitata. 

New records. — Arctic Alaska: Off Point Barrow base, 15 miles 
from shore, 78.2 fms., on bottom of small rocks with worm tubes (1 
station, 1 specimen). Northwest Greenland: Off Conical Rock, 
20-40 fms., 1938; 1 mile northwest Conical Rock, 1940; west side 
Wolstemholm Island, 1940; all collected by Bartlett. East Coast 
North America: Off Nova Scotia, Maine, Massachusetts. Rhode 
Island, low water to 410 fms., U. S. Fish Commission. Alaska: 
Eastern Harbor, Sitka, gravelly-mud, 15 fms., 1874; between Pin- 
nacle and Ulakhta, Unalaska, 16 fms., 1874; Captain's Harbor, 
Unalaska, 25-75 fms., coarse sand, 1874; all collected by Dall. 
Albatross Sta. Herendeen Bay, 1890. Washington: Albatross Sta. 
2876, 48°33' N., 124°53' W., 59 fms., and Sta. 2879, 48°53' N., 125°53' 
W., 34 fms., 1888; Strait of Juan de Fuca, Washington and Puget 
Sounds, low water to 165 fms., Pettibone. 

Distribution. — Widely distributed in the Arctic : Siberian, Alaskan, 
and Canadian Arctic, Baffin Bay, Davis Strait, Greenland, Spits- 
bergen, Norway, Barents Sea, White Sea, Novaya Zemlya, Kara 



274 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m 

Sea. Also Iceland, Faroes to Portugal, Azores, Madeira, Mediter- 
ranean, Adriatic; to Rhode Island; to Gulf of California and Mexico; 
Japan; South Atlantic and Antarctic. In low water to 1,889 fathoms; 
epitokous phase in surface layers. 

Family Goniadidae 

Body elongated, more or less attenuated at both ends, divided 
into two regions, anterior region with uniramous parapodia (fig. 31, 
f) and long, wider posterior region with biramous parapodia (fig. 31, 
g). Segments uniannulate. Prostomium sharply conical, trans- 
versely annulated, ending distally in four small antennae (fig. 31, e). 
Elongated conical dorsal and ventral cirri (fig. 31, /, g). Two anal 
cirri. Notosetae simple; neurosetae compound. Branchiae absent. 
Proboscis eversible, long, cylindrical, with terminal papillae and 
proboscidial organs, a pair of larger, dentate, dark horny macrognaths, 
and a circlet of many smaller H- or Y-shaped micrognaths. Epitoky 
involves only biramous portion. 

Represented by a single species from Point Barrow. 

Genus Glycinde F. Miiller, 1858 
Glycinde wireni Arwidsson, 1898 

FiGUEE 31, e-g 

Goniada nordmanni Wir^n, 1883, p. 403, pi. 30, figs. 4, 5, pi. 32, figs. 1, 2; (not 

Eone nordmanni Malmgren, 1867). 
Glycinde wirini Arwidsson, 1898, p. 53, pi. 3, figs. 48, 49. — not Moore, 1908, p. 

348 ( = (?. armigera Moore; in USNM). 

Description. — Length up to 45 mm., width without setae 2.5 mm., 
width including setae 5 mm., 120 segments. Anterior region rounded, 
tapering sharply to pointed anterior end; posterior two-thirds wider, 
more flattened, tapering gradually posteriorly. Prostomium with 10 
annuli; two pairs of small deeply-set eyes, one pair in basal ring, one 
pair in distal ring (may be obscure). Anterior uniramous region 
consisting of 30-32 (31 in Arwidsson) setigers, with presetal lobe wide 
basally, narrowed abruptly to a lanceolate tip; postsetal lobe wide, 
conical; pre- and postsetal lobes as well as dorsal and ventral cirri 
subequal in length; a fan-shaped bundle of compound, spinigerous 
neurosetae. Posterior biramous region with shorter notopodia dis- 
tinctly separated from neuropodia. Notopodium a short, rounded 
lobe with a short dorsal cirrus and a short, protruding presetal lobe; 
notosetae few in number, acicular, hooked at tip, with long pointed 
hood. Neuropodia with short conical, postsetal lobe, subequal to 
ventral cirrus; presetal lobe wide basally with a lanceolate narrower 
tip, longer than postsetal lobe; neurosetae as in anterior region. 



MARINE POLYCHAETE WORMS — PETTIBONE 275 

except they may be longer and more numerous. Proboscis without 
chevrons; proboscidial organs (horny, yellow, spinous processes) of 
several kinds arranged in longitudinal series. Color: In alcohol: 
Uniformly greenish brown or somewhat banded with one wide, dark- 
greenish-brown band per segment; iridescent. 

New records. — Arctic Alaska: Off Point Barrow base, up to 4 
miles from shore, 23 to 29.1 fms., on various combinations of mud, 
sand, gravel, stones, rocks, shells (3 stations, 5 specimens). 

Distribviion. — Arctic Alaska to Bering Sea. In 23 to 29 fathoms. 

Family Lumbrineridae 

Body elongate, cylindrical, tapered slightly anteriorly and more 
strongly posteriorly, superficially resembling oligochaetes. Pro- 
stomium conical to rounded, with a pair of nuchal organs on posterior 
margin, without antennae or palps (fig. 31, h). First two segments 
apodous and achaetous, without tentacular cirri. Without dorsal 
and ventral cirri. Two to four anal cirri. Parapodia imiramous. 
Proboscis eversible, with dark, chitinous, well-developed masticatory 
apparatus consisting of a pair of ventral mandibles and four pairs of 
maxillary plates (fig. 31, i,j). 

Represented by a single species from Point Barrow. 

G^nus Lumbrineris Blainville, 1828 
Lumbrineris fragilis (O. F. Miiller, 1776) 

Figure 31, h-n 

Lumbricus fragilis Miiller, 1776, p. 216; 1788, p. 22, pi. 22, figs. 1-3. 

Lumbrinereis fragilis Malmgren, 1867, pi. 14, fig. 83. — Chamberlin, 1920, p. 15. — 
Eliason, 1920, p. 33.— Berkeley and Berkeley, 1943, p. 130. 

Lumbriconereis minuta Th^el, 1879, p. 42, pi. 4, figs. 57-59. — Fauvel, 1911, p. 
22.— Augener, 1928, p. 732.— Wesenberg-Lund, 1950a, p. 28; 1950b, p. 71. 

Lumbriconereis fragilis Fauvel, 1923, p. 430, fig. 171, k, I. — Augener, 1928, p. 
730, pi. 11, fig. 5.— Annenkova, 1937, p. 167; 1938, p. 167.— Gorbunov, 1946, 
p. 39.— Thorson, 1946, p. 74.— Zatsepin, 1948, p. 124, pi. 31, fig. 14,— 
Wesenberg-Lund, 1950a, p. 27; 1950b, p. 69; 1951, p. 55. 

Lumbrineris fragilis Hartman, 1944a, pp. 335, 340. 

Description. — Length 150-250 mm., width without setae up to 7 
mm., width with setae 12 mm. Prostomium conical, pointed. First 
two achaetous segments about equal to the following segments or the 
first one slightly longer. Parapodia without branchiae, with presetal 
lobe short, rounded ; postsetal lobe in anterior region of body slightly 
longer, rounded, diagonally truncate; in middle and posterior regions 
of body, postsetal lobe extended somewhat dorsally, somewhat digiti- 
form; with arched limbate setae with fine capillary tips, dark amber- 
colored on basal half, on first 60-100 setigers; with simple (not com- 
pound), hooded, hooked setae, dark amber-colored basally, beginning 



276 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m 

on setigers 15-24 (22-35 or more) ; on young specimens, the hooded, 
hooked setae may begin on more anterior segments; acicula black. 
With two anal cu'ri on ventral side of short, disclike pygidium. Jaw 
formula of proboscis (numbers refer to number of teeth) : M II, 4+4 
(sometimes 5?); M III, 1 + 1 (or 2 + 1); M IV, 1 + 1. Color: In hfe: 
Reddish orange, iridescent. In alcohol: Colorless to gray, iridescent; 
young specimens uniformly reddish brown or somewhat banded with 
one wide dark brown band per segment. 

Remarks. — Three small specimens (0.8 mm. wide without setae) 
have the hooded, hooked setae beginning on the first one to three 
setigers as in L. minuta Theel. They are considered to be the young 
of L. jragilis as has been suggested earlier by Levinsen (1882) and 
Eliason (1920). 

New records. — Arctic Alaska: Off Point Barrow base, up to 12.1 
miles from shore, 20-123.5 fms., on bottoms of mud, stones, worm tubes 
and various combinations of mud, sand, gravel, stones, rocks, shells 
(16 stations, 29 specimens). East Greenland: Off Cape Hold with 
Hope, 23-40 fms., Bartlett, 1939. Eastern North America: Off 
Labrador, 5-13 fms.. Blue Dolphin Expeditions, 1949, 1950; off Nova 
Scotia, Bay of Fundy, Maine, Massachusetts, Rhode Island, 7-452 
fms., U. S. Fish Commission; Woods Hole region, low water and 
dredged, Pettibone, 1950, 1951. Alaska: Chiachi Islands, 20 fms., 
mud, Dall, 1874. 

Distrih'ution. — Widely distributed in the Arctic: Siberian and Alas- 
kan Arctic, Davis Strait, Greenland, Jan Mayen, Spitsbergen, Franz 
Josef Land, Barents Sea, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes, Norway to Azores, Madeira, Mediterranean; Hudson Bay to 
Rhode Island; Bering Sea; north Japan Sea. In low water to 1,883 
fathoms. 

? Larval Lumbrineris fragilis (O. F. Miiller) 

On two occasions nonpelagic larval polychaetes were found in a 
mucous mass off Point Barrow base: September 9, 1948, 21 fathoms, 
from breaking rock (the larvae described as rose-red); September 6, 
1949, 36 fathoms, attached to tunicate Boltenia echinata. Prostomium 



Figure 31. — Glyceridae: a, Glycera capiiata, dorsal view prostomium; b, same, parapodium 
from anterior region, posterior view; c, same, parapodium from middle region, anterior 
view; d, same, one of jaws. Goniadidae: e, Glycinde tvireni, dorsal view anterior end; 
/, same, uniramous parapodium from anterior region, anterior view; g, same, biramous 
parapodium from posterior region, anterior view. Lumbrineridae: A, Lumbrineris 
fragilis, dorsal view anterior end; i, same, maxillary plates of jaw apparatus (one of each 
pair shown); j, same, mandibles of jaw apparatus; k, same, parapodium from anterior 
region, posterior view; /, same, parapodium from posterior region, posterior view; m, 
same, limbate seta with capillary tips; n, same, simple hooded hooked seta (crotchet). 
(For explanation of symbols see p. 210.) 



MARINE POLYCHAETE WORMS — PETTIBONE 

Ir. an -dC "PO^, 



277 




J. ^ - N 

Figure 31. — For explanation see facing page. 



278 PROCEEDINGS OP THE NATIONAL MUSEUM vol. i03 

conical, without appendages. Two wide peristomial segments, the 
second with wide transverse ciliated band. Setigerous segments 8, 
gradually tapering posteriorly, each parapodium with single limbate 
capillary seta and hooded seta. Pygidium bulbous, with wide trans- 
verse ciliated band. 

Family Orbiniidae (Ariciidae) 

Body long, vermiform, with very numerous segments, divided into 
two weakly to sharply separable regions: (1) thoracic: More or less 
flattened dorsoventrally and enlarged; attenuated anteriorly; neuro- 
podia cushionlike, with several rows of setae; notopodia more or less 
cirriform, a bundle of crenulate capillary setae (fig. 32, 6); (2) ab- 
dominal: Much longer, semicylindrical; neuropodia bilobed, project- 
ing dorsally (fig. 32, c). Branchiae medial to notopodia, simple, 
dorsal, erect, lanceolate or straplike, strongly ciliated, a pair on aU 
segments except the anterior ones. Prostomium conical or globular, 
without appendages, with or without two small eyes, with two ciliated 
nuchal organs (fig. 32, a). First one or two segments apodous and 
achaetous. Usually without dorsal or ventral cirri. Setae simple, of 
varied form. Pygidium with two to four anal cirri. Proboscis 
eversible, unarmed, a voluminous, weakly- to much-branched, soft sac. 

Represented by a single species from Point Barrow. 

Genus Scoloplos Blainville, 1828 
Scoloplos armiger (O. F. Miiller, 1776) 

Figure 32, a-e 

Lumhricus armiger O. F. MuUer, 1776, p. 215; 1788, p. 22, pi. 22, figs. 4, 5. 

Aricia arctica Hansen, 1882, p. 34, pi. 5, figs. 20-26. — Murdoch, 1885, p. 154. 

Scoloplos armiger Fauvel, 1914, p. 224; 1927, p. 20, fig. 6, k-g. — Eliason, 1920, 
p. 39. — Augener, 1928, p. 742.— Monro, 1930, p. 145. — Annenkova, 1931, p. 
203; 1937, p. 169; 1938, p. 170.— Okuda, 1937a, p. 102, fig. 4.— Wesenberg- 
Lund, 1939b, p. 12; 1950a, p. 29; 1950b, p. 73; 1951, p. 58.— Berkeley and 
Berkeley, 1942, p. 195; 1952, p. 97, figs. 197-199.— Hartman, 1944a, pp. 336, 
340, pi. 18, fig. 5 (not pi. 19, fig. 6 (=Orbinia)) .—Thorson, 1946, p. 78, fig. 
37.— Gorbunov, 1946, p. 39.— Zatsepin, 1948, p. 128, pi. 32, fig. 1. 

Haploscoloplos alaskensis Hartman, 1948, p. 30, fig. 8, a-c. 

Description. — ^Length up to 120 mm., width 2 mm. Prostomium 
conical, pointed, with two deeply buried eyes (not visible when pre- 
served). First segment achaetous and apodous. Branchiae first 
appear on setigers 12 or 13 (9-17), very small at first, then triangular, 
then ligulate. Thoracic region consisting of about 17 setigers 
(12-20) ; notopodia with conical postsetal lobes and numerous, finely 
crenulate setae with capillary tips; neuropodia elongate, cushionlike, 
with a median postsetal conical lobe, with numerous rows of crenulate 



MARINE POLYCHAETE WORMS — PETTIBONE 279 

setae with capillary tips; some of these setae in the lower part of the 
neuropodial bundle may be worn down, with blunt tips resembling 
crotchets. These so-called crotchets may be more or less numerous 
on all thoracic segments or on some only, or may be lacking. Ab- 
dominal region having an exceedingly straggly appearance with the 
dorsally directed branchiae and parapodial lobes; notopodia with 
erect, digitiform postsetal lobes and capillary crenulate setae and 
sometimes also with forked setae; neuropodia with two conical, 
unequal lips, with capillary crenulate setae and an elongate thickened 
parapodial flange below the neuropodial lobe. On the last few thoracic 
and first few abdominal segments there may be one or two extra 
conical podal lobes, one at the level of the lower part of the neuropodial 
setae and one subpodal; they are in the transitional region of thoracic 
and abdominal regions and become part of the parapodial flanges of 
the abdominal segments. Without intermediate interramal cirri, 
but with lateral, ciliated organs between the rami. Pygidium with 
two filiform anal cirri. Proboscis soft, more or less lobed. Color: 
In alcohol: Without color or yellowish brown, with darker areas on 
the elongate neuropodial flanges of the abdominal region and on the 
branchiae. 

Remarks. — Haploscoloplos alaskensis Hartman from southwestern 
Alaska is herein referred to S. armiger. It differs from the typical 
S. armiger in lacking crotchets in the thoracic neuropodia. As 
indicated previously by Fauvel (1914) and others, this seems to be a 
variable character, perhaps associated with the substratum and age. 
The crotchets were not present on small specimens from Point Barrow, 
and they may be more numerous in some parapodia than in others 
and more abundant in the lower part of the neuropodia; they appear 
to be worn crenulate capillary setae. On certain substrata, pre- 
sumably, the setae may not be worn and thus the so-called crotchets 
are absent. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow; west side Elson Lagoon near entrance to small lagoon 
to the west, 1.2 fms.; off Point Barrow base, up to 12.1 mQes from 
shore, 1.7-75.5 fms., on bottoms of mud, stones, and various com- 
binations of mud, gravel, stones, rocks, large perforated rocks (12 
stations, 19 specimens). Between Icy Cape and Cape Lisbume, 
15-20 fms., mud, sand, Dall, 1874. East Coast North America: 
Bay of Fundy, Maine, Massachusetts, 13—48 fms., U. S. Fish 
Commission. 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Davis Strait, Greenland, Jan Mayen, Spitsbergen, 
Franz Josef Land, White Sea, Novaya Zemlya, Kara Sea. Also 
Iceland, Faroes, Shetlands to France, Mediterranean; Labrador to 



280 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Massachusetts; southern Alaska to British Columbia; Japan; South 
Pacific (Chile); Antarctic; off South Africa. In low water to 1,100 
fathoms. 

Family Spionidae 

Body vermiform, without distinct regions. Prostomium small, 
elongated, scarcely separated from an enlarged peristomium lateral to 
prostomium, with or without eyes (usually four), antennae usually 
lacking (may be a small occipital antenna), two very long, extensile, 
tentaculiform, longitudinally grooved, easily deciduous palps (fig. 
32, /, i, I, r) . Parapodia biramous, with dorsal and ventral postsetal 
lamellae more or less developed; dorsal and ventral cirri lacking 
(fig. 32, g, s). With ligulate or cirriform dorsal branchiae on upper 
part of parapodial bases on a certain number of segments (rarely 
pinnate, rarely lacking). Setae simple capillaries and hooded 
crotchets (fig. 32, h, k, q). Pygidium terminated in an anal cup 
(fig. 32, p) or anal cirri. Proboscis short, scarcely or somewhat 
protrusible, may be somewhat lobulated. 

Represented by four genera and four species from Point Barrow. 
All genera have branchiae present on certain segments and prostomia 
without frontal horns. 

Key to the genera of Spionidae from Point Barrow 

1. Setigerous segment 5 strongly modified with special large amber-colored 

setae and without dorsal and ventral lamellae (fig. 32, m, n). Branchiae 
numerous, paired, dorsal, simple, straplike. Notopodia with capillary 
setae only; neuropodia with capillary setae and hooded bidentate crotchets 
beginning on setigers 7 or 8 (fig. 32, q). Pygidium ending in anal cup 

(fig. 32, p) Polydora (p. 280) 

Setigerous segment 5 not modified 2 

2. Branchiae on only few anterior segments, 3-11 pairs, often pinnate (fig. 

32, j), beginning on setiger 2. Capillary setae and hooded crotchets in both 

noto- and neuropodia. Pygidium with anal cirri Prionospio (p. 282) 

Branchiae more numerous, simple, straplike. Notopodia with capillary setae 
only; neuropodia with capillary setae and hooded crotchets 3 

3. Branchiae begin on setiger 1, continuing up to last setigers. Pygidium with 

anal cirri Spio (p. 284) 

Branchiae begin on setiger 2, continuing on large number of segments. Py- 
gidium with anal cup, may be somewhat lobulated Nerinides (p. 285) 

Genus Polydora Rose, 1801 
Polydora caulleryi Mesril, 1897 

Figure 32, l-q 

Polydora caulleryi Mesnil, 1897, p. 88, pi. 3, figs. 12-16.— Fauvel, 1927, p. 54, 

fig. 19, f-h. 
Polydora brachycephala Hartman, 1936c, p. 48, figs. 3-5; 1944b, p. 258. 



MARINE POLYCHAETE WORMS — PETTIBONE 281 

Description. — Up to 50 mm. long, 2.2 mm. wide. Prostomium 
feebly notched anteriorly or distinctly bifid, prolonged posteriorly in 
a crest extending to setiger 4 (to 6) , without distinct occipital antenna, 
with fom- eyes in trapezoidal arrangement (or six in young specimens) 
or eyes lacking. Palps usually missing in alcohol. A protrusible, 
somewhat lobulated, brownish proboscidial region. Branchiae begin 
on setigers 7 or 8, continuing on numerous segments, absent from 
more posterior segments; branchiae short (about length of capillary 
notosetae), straplike, continuous with dorsal lamellae. Modified seti- 
ger 5 with group of capillary notosetae, usually 5-7 (3-12) special 
amber-colored hooks, and group of capillary neurosetae; special hooks 
all similar, falcate, with pectinate top. With conspicuous, somewhat 
conical dorsal and. ventral postsetal lamellae on setigers 1-4, lacking 
on modified setiger 5, becoming rather low and wide on more posterior 
segments. With capillary setae in both notopodia and neuropodia; 
beginning on setiger 7, neuropodial capillary setae mostly replaced 
by hooded bidentate crotchets. In more posterior setigers (last 20 
or so), notopodia conical, lacking branchiae or lamellae, with conical 
bundle of awl-shaped setae scarcely surpassing notopodial tip and 
small bundle of capillary setae; neuropodia with two or three crotchets 
and few capUlary setae. Anal disc with complete middorsal notch, 
shorter midventral and midlateral notches, thus 4-lobed. Color: In 
alcohol: Without color or grayish. 

Remarks. — The type of P. brachycephala Hartman from central 
California was examined and is herein referred to P. caulleryi. 

New records. — Arctic Alaska: Off Point Barrow base, up to 8 
miles from shore, 16.7-75.5 fms., on bottoms of stones and various 
combinations of mud, gravel, stones, rocks, large perforated rocks, 
with bryozoans and hydroids (8 stations, 16 specimens). West Coast 
North America: Washington and Puget Sounds, low water to 50 
fms., Pettibone. 

Distribution. — Arctic Alaska to central California; Ireland, Great 
Britain to France. In low water to 75.5 fathoms. 

Polydora larva 

A larva of a Polydora was taken in plankton August 2, 1948. This 
was perhaps the larva of P. caulleryi Mesnil since it was the only 
species of Polydora found in the Point Barrow region. Prostomium 
rounded, with two palps, and with black chroma tophores. Fifth 
setigerous segment modified, with short, thick, slightly curved acicular 
setae; rest of 21 setigerous segments with long larval capillary setae. 
Pygidium a rounded disc, notched middorsaUy, with black chromato- 
phores. Black chroma tophores arranged transversely on setigers 2-5, 
then in two longitudinal lines on each side of middorsal line, extending 
to anal cup. 



282 PROCEEDINGS OF THE NATIONAL MUSEUM tol.io3 

C^nus Prionospio Malmgren, 1867 
Prionospio malmgreni Clapar^de, 1868 

Figure 32, i-k 

Prionospio malmgreni ClaparMe, 1868, p. 333, pi. 22, fig. 3 (fide Fauvel, 1927). — 
Ehlers, 1913, p. 511.— Fauvel, 1927, p. 61, fig. 21, a-e.— Annenkova, 1937, 
p. 171; 1938, p. 174.— Hartman, 1948, p. 36.— Wesenberg-Lund, 1951, 
p. 70. — Berkeley and Berkeley, 1952, p. 29, figs. 54, 55. 

Spiophanes tenuis Verrill, 1879b, p. 176. 

Prionospio tenuis Verrill, 1882, p. 370. — Hartman, 1944a, pp. 336, 340, ? pi. 19, 
fig. 7; not 1945, p. 32 ( = P. treadwelli Hartman, 1951). 

Description. — Length up to 25 mm., width 1 mm. Prostomium 
dilated anteriorly, with anterior border rounded, narrowing posteriorly, 
extending in obtuse crest to setiger 2 (to 4) . Four eyes, two smaller, 
two much larger (may be lacking) . Two long palps, usually missing 
(in alcohol). A short, eversible, somewhat lobulated proboscidial 
region. Branchiae four pairs, on setigers 2-5, completely separated 
from dorsal lamellae, pairs 1 and 4 longer, pinnate, somewhat decid- 
uous, pairs 2 and 3 shorter, smooth, triangular. Dorsal postsetal 
lamellae of first setiger small, much larger and triangular on next 
four setigers, then diminishing in size, becoming reduced beginning 
on setigers 8 or 9; ventral postsetal lamellae oval, rounded. Hooded, 
multidentate crotchets beginning about setigers 12-16 in neuropodia 
and about setiger 40 in notopodia. Beginning on setiger 7, slightly 
elevated transverse folds between dorsal lamellae, continuing on next 
6-14 segments. Pygidium with longer, unpaired dorsal cirrus and 
two smaller lateral ones. Without color in alcohol. 

Remarks. — The types of Prionospio tenuis Verrill from Cape Cod 
Bay were examined and are herein referred to P. malmgreni. There 
are four pairs of branchiae, the first and fourth pairs pinnate, the sec- 
ond and third pairs shorter, smooth. 

New records, — Arctic Alaska: Off Point Barrow base, 4 miles from 
shore, 29 fms., on bottom of gravel, small stones (1 station, 2 speci- 



FiGURE 32. — Orbiniidae: a, Scoloplos armiger, dorsal view anterior end; i, same, parapodium 
from thoracic region, posterior view; c, same, parapodium from abdominal region, anterior 
view; d, ty same, crenulate neurosetae from thoracic region. Spionidae:/, Spio filicornis, 
dorsal view anterior end (palps missing); g, same, parapodium from anterior region, 
anterior view; h, same, hooded bidentate crotchet; i, Prionospio malmgreni, dorsal view 
anterior end; /, same, branchiae on setigers 2-S (one of each pair shown); k, same, hooded 
multidentate crotchet; /, Polydora caulleryi, dorsal view anterior end (palps missing); 
m, same, parapodium of modified fifth setigerous segment; «, same, special hooks from 
setiger 5; o, same, notopodium of posterior setiger, with conical bundle awl-shaped 
setae; p, same, anal disc of pygidium, lateral view; q, same, hooded bidentate crotchet, 
f, Ntrinides sp., dorsal view anterior end; s, same, parapodium from anterior region. 
(For explanation of symbols, see p. 210.) 



MARINE POLYCHAETE WORMS — PETTIBONE 283 




P Q 

Figure 32. — For explanation see facing page. 



261112—54 6 



284 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

mens). West Coast North America: Puget Sound, Washington, 
low water, mud, Pettibone. East Coast North America: Off 
Labrador, 40 fms,, Blue Dolphin Expedition, 1949; off New Hamp- 
shire and Massachusetts, 25-48 fms., U. S. Fish Commission. 

Distribution. — Arctic Alaska to Washington; north Japan Sea; 
Iceland, Norway to Mediterranean; Labrador to Massachusetts; 
South Africa. In low water to 250 fathoms; surface. 

Genus Spio Fabricius, 1785 
Spiofilicornis (Miiller, 1776) 

Figure 32, f-h 

Nereis filicornis Miiller, 1776, p. 218. 

Spio filicornis Malmgren, 1867, p. 91, pi. 1, fig. 1. — Eliason, 1920, p. 40, figs. 
7-9.— Fauvel, 1927, p. 43, fig. 15, a-g.— Augener, 1928, p. 738, pi. 11, fig. 7.— 
Annenkova, 1937, p. 169; 1938, p. 173.— Hartman, 1941a, p. 293; 1948, p. 
36.— Berkeley and Berkeley, 1943, p. 130.— Thorson, 1946, p. 93, fig. 46.— 
Zatsepin, 1948, p. 132, pi. 32, fig. 11.— Wesenberg-Lund, 1950a, p. 30; 
1950b, p. 76; 1951, p. 71. 

Spio limicola Verrill, 1879, p. 176. 

Spio filicornis var. pacifica Berkeley and Berkeley, 1936, p. 475; 1952, p. 25, figs. 
47, 48. 

Description. — Up to 30 mm. long, 2 mm. wide. Prostomium bell- 
shaped to conical, widest anteriorly, rounded, tapered posteriorly to 
short crest extending on first two setigers. Four eyes in trapezoidal 
arrangement, anterior pair larger. Two palps missing (in alcohol). 
A short, eversible, somewhat lobulated proboscidial region. Branchiae 
begin on setiger 1, continuing to posterior end, simple, straplike. 
Dorsal postsetal lamellae of more anterior segments short, wide, with 
branchiae fused to upper sides; in more posterior segments, lamellae 
cirriform, separate from branchiae. Ventral postsetal lamellae 
rounded, not notched. Capillary setae only in notopodia. Neuropodia 
with hooded, bidentate crotchets beginning on setigers 9-18, about six 
in number (5-10). Pygidium with four cirri? Color: In alcohol: 
Yellowish white with dark brown pigment on peristomial region and 
along each side of anterior fourth of body. 

Remarks. — The types of Spio limicola Verrill from Cape Cod Bay 
were examined and are herein referred to S. jilicornis. S. limicola 
differs from S. setosa Verrill, to which it has been referred (Hart- 
man, 1942, p. 63), in the number of neuropodial hooded crotchets, 
6-9 in S. limicola and about 16 in ^S. setosa. 

New records. — Arctic Alaska: Off Point Barrow base, up to 8 
miles from shore, 36-75.5 fms., on various combinations of gravel, 
rocks, stones, large perforated rocks (2 stations, 2 specimens) . West 
Coast North America: Washington Sound, low water, Pettibone. 
East Coast North America: Off Massachusetts, 15-31 fms., U. S. 
Fish Commission. 



MARINE POLYCHAETE WORMS — PETTIBONE 285 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Davis Strait, Greenland, Spitsbergen, Novaya Zemlya. 
Also Iceland, Faroes, Norway to France; Hudson Bay to Massachu- 
setts; Bering Sea to central California; Japan. In low water to 211 
fathoms; sm-face. 

Genus Nerinides Mesnil, 1896 
Nerinides sp. 

Figure 32, r, s 

Two small specimens obtained from plankton 10 feet from shore, 
agreeing in some respects with the description of N. cantabra Rioja 
(Fauvel, 1927, p. 31). Length 16 mm., width 1 mm., segments 57. 
Prostomium rounded anteriorly, tapering posteriorly, with middorsal 
crest extending on first setiger, with four eyes, one pair larger, a 
raised area just posterior to eyes (median antenna?), with two large, 
thick palps (fig. 32, r). Dorsal and ventral postsetal lamellae distinct, 
dorsal ones may be slightly undulate, ventral ones entire, not notched 
(fig. 32, s). Branchiae begin on setiger 2, continuing to near posterior 
end, fused to upper part of dorsal lamellae. Body with low transverse 
dorsal ridges between bases of branchiae. First setiger with both 
dorsal and ventral setae. Notopodia with only capillary setae, being 
extra long in posterior region. Neuropodia with capillary setae and 
hooded bidentate crotchets beginning around setiger 20. Pygidium 
with 4-lobed membranous appendage. Without color in alcohol. 

Family Cirratulidae 

Body more or less cylindrical, attenuated at both ends, with 
numerous, short, compact, nearly similar segments. Prostomium 
small, usually without appendages, with or without eyes. Pair of large 
grooved spioniform palps (fig. 33, d) or with more or less numerous 
grooved tentacular fiilaments inserted on dorsal surface of an anterior 
segment (fig. 33, a). Branchiae simple, filamentous, long, con- 
tractile, inserted above notopodia on few to many segments (may 
be absent). Parapodia biramous, lobes scarcely projecting (fig. 33, 
6). Setae usually simple, some capillary, others acicular crotchets 
(fig. 33, c) . Without dorsal, ventral, or anal cirri. Proboscis smooth, 
unarmed. May have epitokous pelagic form at time of reproduction. 
Usually in mud; some (Dodecaceria) in calcareous matrix. 

Represented by two genera and two species from Point Barrow. 
Both genera have prostomium conical, buccal segment and two fol- 
lowing segments achaetous, more or less fused to prostomium, with 
lateral filamentous branchiae beginning just anterior to first setigerous 
segment, continuing on large number of segments. 



286 PROCEEDINGS OF THE NATIONAL MUSEUM vol.108 

Key to the genera of Cirratulidae from Point Barrow 

1. With few to numerous, grooved tentacular filaments just anterior to first 
setigerous segment, subequal to branchiae (without 2 palps; fig. 33, a). 

Cirratulus (p. 286) 

With 2 large grooved, spioniform palps just anterior to first setigerous segment 

(without tentacular filaments; palps deciduous; may be missing in epitoke; 

fig. 33, d) Chaetozone (p. 287) 

Genus Cirratulus Laxuark, 1801 
Cirratulus cirratus (Miiller, 1776) 

Figure 33, a-c 

Lumbricus cirratus Miiller, 1776, p. 215. 

Cirratulus cirratus Malmgren, 1867, p. 95. — Chamberlin, 1920, p. 20. — Eliason, 
1920, p. 57.— Fauvel, 1927, p. 94, fig. 33, a-g; 1933, p. 46; 1936, p. 72.— 
Augener, 1928, p. 750, pi. 11, fig. 9.— Monro, 1930, p. 154; 1936, p. 161.— 
Annenkova, 1931, p. 203; 1932, p. 136; 1934, p. 322; 1937, p. 173; 1938, p. 
181.— Berkeley and Berkeley, 1942, p. 197; 1943, p. 130; 1952, p. 31, figs. 
58, 59.— Hartman, 1944a, pp. 334, 341; 1944b, p. 263; 1948, p. 37.— Gorbunov, 
1946, p. 39.— Zatsepin, 1948, p. 133, pi. 32, fig. 18.— Wesenberg-Lund, 
1950a, p. 33; 1950b, p. 81; 1951, p. 74.— Hartman and Reish, 1950, p. 34. 

Cirratulus robustus Johnson, 1901, p. 423, pi. 14, figs. 149, 150. — Rioja, 1941, 
p. 728.— Berkeley and Berkeley, 1942, p. 197. 

Cirratulus cingulatus Johnson, 1901, p. 422, pi. 14, figs. 145-148. — Rioja, 1941, 
p. 729. 

Cirratulus spectabilis Berkeley and Berkeley, 1952, p. 32. 

Description. — Length up to 300 ^mm., width 5 mm. Prostomium 
variable in shape: Short, broad, bluntly conical; almost square- 
shaped; or tapered gradually, then abruptly. Prostomium with 
usually four or five (2-8) eyes on each side in obliquely transverse 
row or may be united in almost solid arc. Buccal segment more or 
less fused with prostomium, two following achaetous segments more 
or less distinct. Tentacular filaments anterior to first setigerous 
segment, in two groups of 2-24 in obliquely transverse rows, long, 
filamentous, grooved longitudinally. Branchiae begin just anterior to 
first setigerous segment, lateral to group of tentacular filaments, 
continuing through greater part of body, anteriorly inserted close to 
upper part of notopodia, then gradually shifting more dorsally and 
then sUghtly more ventrally. Parapodia consisting of slightly pro- 
jecting tori common to both no to- and neuropodia, with capillary 
setae in both lobes, with additional acicular crotchets first appearing 
in neuropodia and more posteriorly in notopodia also. Anus sub- 
dorsal. Color: In life: Orange yellow with red branchiae; dark 
orange brown with orange-red branchiae. In alcohol: Yellowish 
brown or without color. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 5-6.6 fms.; off Point Barrow base, up to 8 miles from 



MARINE POLYCHAETE WORMS — PETTIBONE 287 

shore, 13.3-75.5 fms., on bottoms of stones, and various combinations 
of mud, pebbles, rock, stones, gravel, large perforated rocks, with 
bryozoans, embedded in mud at bases of barnacles and bryozoans as 
Eucratea loricata (19 stations, 183 specimens). Siberia: Plover Bay, 
10-35 fms., Dall, 1880. Bering Sea: Albatross station, Nikolski, 
Bering Island, 1892; St. George Island, Pribilofs, 2 miles off shore, 
40 fms., Hanna, 1913; Atka Island, Aleutians, L. M. Turner, 1879. 
West Coast North America: Strait of Juan de Fuca, Washington 
and Puget Sounds, low water, Pettibone. East Coast North Amer- 
ica: Off Labrador, 95 fms.. Blue Dolphin Expedition, 1949; oil Maine, 
Massachusetts, low water to 67 fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Barents Sea, Novaya 
Zemlya. Also Iceland, Faroes, Norway to France, Canary Islands; 
Hudson Bay to Massachusetts; Bering Sea to Mexico; north Japan 
Sea to Japan, Manchuria. Southern latitudes: Falkland Islands, 
Magellan Straits, South Georgia, Kerguelen. In low water to 1,611 
fathoms. 

Genus Chaetozone Malmgren, 1867 

Chaetozone setosa Malmgren, 1867 

Figure 33, d 

Chaetozone setosa Malmgren, 1867, p. 96, pi. 14, fig. 84. — Th^el, 1879, p. 54, pi. 
4, figs. 49-51.— Fauvel, 1914, p. 217, pi. 20, fig. 1; 1927, p. 101, fig. 35; 
1934a, p. 47.— Eliason, 1920, p. 57. — Augener, 1928, p. 750. — Gustafson, 
1936, p. 8.— Monro, 1937, p. 301.— Annenkova, 1937, p. 174; 1938, p. 182.— 
Berkeley and Berkeley, 1942, p. 197; 1952, p. 35, fig. 63.— Hartman, 1944a, 
pp. 334, 341.— Zatsepin, 1948, p. 133, pi. 32, fig. 16.— Wesenberg-Lund, 
1950a, p. 34; 1950b, p. 81; 1951, p. 75. 

Description. — Length up to 25 mm., width 2 mm. Prostomium 
conical, acutely pointed, without eyes. Buccal segment and two 
following achaetous segments more or less fused; in specimens from 
Point Barrow, buccal segment fused with prostomium, second and 
third segments fused dorsally. Paired palps large, long, grooved, 
inserted just anterior to first setigerous segment, deciduous, often 
missing on preserved specimens and missing when in epitokous phase. 
Branchial filaments begin just anterior to first setigerous segment, 
immediately posterior to palps, continuing on large number of anterior 
segments, lacking on posterior region, readily deciduous, inserted just 
posterior to upper part of parapodial tori. Parapodia are slightly 
projecting tori common to both noto- and neuropodia. In anterior 
region, both lobes with long, delicate capillary setae; more posteriorly, 
notopodia with still longer capillary setae, neuropodia with shorter, 
stouter capillary setae and short, unidentate acicular crotchets; more 
posteriorly, with acicular crotchets in both noto- and neuropodia. 



288 PROCEEDINGS OF THE NATIONAL MUSEUM TOL.108 

In epitokous phase, dorsal setae in anterior region much longer; ia 
posterior region, parapodial tori more projecting, dorsal and ventral 
acicular crotchets forming semicircular, fan-shaped group, almost en- 
circHng body. Pygidium with small, rounded to conical lobe, ventral 
and posterior to anus (like a cover for anal opening). Color: In 
alcohol: Light to dark gray, brownish, or iridescent bluish black. 

New records. — Arctic Alaska: Off Point Barrow base, up to 12.1 
miles from shore, 20-123.5 fms., on various combinations of mud, 
stones, gravel, rocks, large perforated rocks, mass of worm tubes, on 
surface of tunicates (12 stations, 29 specimens). East Coast North 
America: Off Labrador, 95-125 fms.. Blue Dolphin Expedition, 1949; 
off Maine, New Hampshire, Massachusetts, 11-58 fms., U. S. Fish 
Commission. 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Davis Strait, Greenland, Spitsbergen, Franz Josef 
Land, Barents Sea, Novaya Zemlya. Also Iceland, Faroes, Norway 
to Canary Islands, Mediterranean, Adriatic, Gulf of Aden; Labrador 
to Massachusetts; Bering Sea to British Columbia; north Japan Sea. 
Southern latitudes: Falkland Islands, Magellan Straits, Kerguelen. 
In low water to 1,333 fathoms; surface. 

Family Flabelligeridae (Chloraemidae) 

Body subcylindrical or subfusiform, with short similar segments 
covered with papillae, often with adherent sand grains, may be 
encased in thick mucous mantle. Prostomium and buccal segment 
fused, with two thick grooved palps, two groups of filiform cephalic 
branchiae, and more or less developed buccal tube around mouth, 
all of which may be partially or completely retracted within anterior 
setigers, with or without setae of anterior setigers directed anteriorly 
forming a cephaUc cage. Parapodia biramous; notosetae simple, 
capillary, ringed; neurosetae simple or pseudocompound, capillary 
or hooked. Without dorsal, ventral, or anal cirri. Anus terminal. 
Usually in muddy sand. 

Represented by two genera and three species from Point Barrow. 

Key to the genera of Flabelligeridae from Point Barrow 

1. Body covered with long pedunculate papillae (fig. 33, g), usually encased in 
thick mucous mantle. Fused prostomium and buccal segment partially 
retractile within first setigers and more or less hidden by cephalic cage formed 
by setae of first setiger, the setae long, directed anteriorly, almost completely 

encircling segment Flabelligera (p. 289) 

Body covered with short papillae, usually encrusted with sand, not encased 
in mucous mantle. Fused prostomium and buccal segment completely 
retractile within anterior setigers, may be completely hidden; without 
cephalic cage formed by anterior setigers; when buccal segment, etc. are 
retracted, anterior end is trilobed (fig. 33, h) Brada (p. 290) 



MARINE POLYCHAETE WORMS — PETTIBONE 289 

Genus Flabelligera Sars, 1829 

Flabelligera affinis Sars, 1829 
Figure 33, e~g 

Flabelligera affinis Sars, 1829, p. 31, pi. 3, fig. 16 (fide Malmgren, 1867). — Malmgren, 
1867, p. 83.— Chamberlin, 1920, p. 21.— Eliason, 1920, p. 62.— Fauvel, 1927, 
p. 113, fig. 40, a-f.— Augener, 1928, p. 768.— Monro, 1930, p. 160, fig. 63; 
1936, p. 164.— Annenkova, 1937, p. 174; 1938, p. 184.— Berkeley and Berkeley, 
1943, p. 130.— Hartman, 1944a, pp. 335, 341, pi. 33, figs. 12, 13 (as ?Brada 
setosa Verrill). — Gorbunov, 1946, p. 39. — Zatsepin, 1948, p. 134, pi. 33, fig. 
1.— St0p-Bowitz, 1948a, p. 8, fig. 1; 1948b, p. 30.— Wesenberg-Lund, 1950a, 
p. 35; 1950b, p. 82; 1951, p. 76. 

Flabelligera infundibularis Johnson, 1901, p. 417, pi. 12, figs. 124-127. — Hartman, 
1938c, p. 14; 1948, p. 40.— Berkeley and Berkeley, 1952, p. 7, figs. 1-4. 

Flabelligera infundibularum Moore, 1923, p. 223. 

Flabelligera infundibuliformis Hartman and Reish, 1950, p. 35. 

Description. — Length up to 130 mm., width including mantle 12-15 
mm., segments up to 7 1 . Body rounded, attenuated shghtly anteriorly 
and more so posteriorly. Body covered with numerous papillae 
which are elongate, thin, flexible, usually somewhat coiled, with larger 
bulbous tips quite variable in shape. Body usually covered with 
thick, mucous, transparent mantle (may be missing in smaU speci- 
mens or specimens appearing in surface waters) which is penetrated 
by the pedunculate papillae. Fused prostomium and buccal segment 
retractile within anterior setiger, with pair of stout, grooved palps, 
two groups of numerous filiform branchiae, two pairs of eyes (almost 
coalesced), and somewhat protruding buccal siphon around ventral 
mouth opening. First setigerous segment with wide, funneUike 
depression into which buccal siphon, etc., can mostly be withdrawn; 
with circlet of elongated, ringed setae around margin and directed 
forward or spread fimnellike, forming cephalic cage; with numerous 
elongate papillae having slender straight peduncles and fusiform tips. 
Notopodia conical, diagonally truncate lobes, with capillary ringed 
setae, with elongate papillae on posterior side that may be up to three- 
fourths length of notosetae (similar in shape to those around cephalic 
cage; different from those on body). Neuropodia conical, each with 
usually one (sometimes two) large, hooked, amber-colored pseudo- 
compoimd seta, ringed basaUy, hooked tips dark to black. When 
mucous mantle is present, setae almost completely embedded and the 
hooked tips of neurosetae and capillary tips of notosetae only ex- 
tending beyond mantle. Pygidium a fleshy, rounded rim; anus 
subdorsal. Color: In alcohol: Without color or slightly reddish 
brown. 

New records. — Arctic Alaska: West side Elson Lagoon near en- 
trance to small lagoon to west, 1.2 fms.; Point Barrow base, washed 
ashore; off Point Barrow base, 18.3-123.5 fms., on various combina- 



290 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

tions of mud, stones, gravel, rocks, with mass of worm tubes (12 
stations, 15 specimens). Bering Sea: Albatross Sta. 3274, 55°34' 
N., 162°31' W., 19 fms., and station at Unalaska, Aleutians, 1890; 
IluUiuk Harbor, Unalaska, Dall, 1871 . West Coast North America : 
Washington and Puget Sounds, 10 fms. and surface, Pettibone. 
Canadian Arctic: Dobbin Bay, east Ellesmere Land, 79°36' N., 
73°35' W., 16 fms., W. H. Littlewood, 1950. Foxe Basin, Baffin 
Island, 66°83' N., 80°07' W., Bartlett, 1927. West Greenland: 
Between Capes Alexander and Chalon, 25-40 fms., Bartlett, 1937. 
East Coast North America: Off Labrador, 10-30 fms., Blue 
Dolphin Expeditions, 1949, 1950; Grand Manan, Maine, New 
Hampshire, Massachusetts, 3-90 fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Franz Josef Land, 
Barents Sea, White Sea, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes, Norway to France, Mediterranean; Hudson Bay to Mas- 
sachusetts; Bering Sea to southern California; north Japan Sea. 
Southern latitudes: Falkland Islands, Magellan Straits, South 
Georgia, South Africa, Australia. In low water to 889 fathoms; 
surface. 

Genus Brada Stimpson, 1854 

Key to the species of Brada from Point Barrow 

1. Body covered with filiform papillae, often encrusted with sand grains, with 
tips of papillae bare. Notopodia distinct, with 2-5 capillary setae per lobe. 

B. villosa 
Body covered with low globular papillae, often encrusted with a thin layer of 
sand grains. Notopodia indistinct, with 0-2 very delicate capillary setae. 

B. inhabilis 

Brada villosa (Rathke, 1843) 

Siphonostoma villosum Rathke, 1843, p. 215, pi. 11, figs. 11, 12. 

Brada villosa Malmgren, 1867, p. 84.— Eliason, 1920, p. 62.— Chamberlin, 1920, 

p. 22.— Fauvel, 1927, p. 121, fig. 43, e-1; 1934a, p. 49.— Monro, 1930, p. 161; 

1937, p. 303.— Annenkova, 1937, p. 175; 1938, p. 185.— Okuda, 1937b, 

p. 53, pi. 2, fig. D.— Berkeley and Berkeley, 1943, p. 130; 1952, p. 7, fig. 5.— 

Zatsepin, 1948, p. 136, pi. 33, fig. 4.— St0p-Bowitz, 1948a, p. 33, fig. 9; 

1948b, p. 39, fig. 15. — Hartman and Reish, 1950, p. 35. — Wesenberg-Lund, 

1950a, p. 35; 1950b, p. 85; 1951, p. 78, fig. 7. 
Brada setosa Verrill, 1873, pp. 431, 434, 508, 606; 1881, p. 302, pi. 9, fig. 4.— 

Sumner, Osburn and Cole, 1913, p. 630. 
Trophonia rugosa Hansen, 1882, p. 38, pi. 7, figs. 9-12. 
Trophonia arctica Hansen, 1882, p. 39, pi. 7, figs. 17-20. 
Brada granulata Murdoch, 1885, p. 155 (not B. granulata Malmgren, 1867). 
Brada pilosa Moore, 1906b, p. 231, pi. 10, figs. 14-17. 
Stylaroides pluribranchiata Moore, 1923, p. 222. 
Brada rugosa St0p-Bowitz, 1948a, p. 37, fig. 10; 1948b, p. 41. 



MARINE POLYCHAETE WORMS — PETTIBONE 291 

Description. — Length 28-60 mm., width 5-9 mm., setigers 23-35. 
Body fusiform, slightly flattened ventrally, convex dorsally, attenu- 
ated and more or less elongated posteriorly, having a characteristic 
rough, hirsute aspect, covered with cylindrical, fusiform to claviform 
papillae, longer dorsally than ventrally, usually encrusted with sand 
grains more basally, with distal tips bare; there are all gradations: 
Without encrusting sand (looks "hairy"), with scattered sand grains, 
and with rather thick layer of sand grains, usually fewer on antero- 
ventral part. Underlying surface nodular and, in larger specimens, 
outer papillate layer may be worn off, the surface appearing rugose, 
with low mounds. Retractile prostomium and buccal region with two 
groups of numerous filiform branchiae borne on pair of dorsolateral 
bosses, pair of thick grooved palps, and ventral eversible buccal siphon 
around mouth. Setae of first setigerous segment 8-10 per bundle, 
slightly more elongate than following, directed anteriorly. Notopodia 
and neuropodia distinct, with long fusiform papillae grouped around 
setae. Notosetae long, capillary, ringed, 2-5 in number. Neurosetae 
beginning on setiger 2, large, 3-6 in number, amber-colored, ending in 
sharp, transparent, and fragile tips. Pair of nephridial papillae near 
anterior border of ventral side of setiger 5. Color: In life and in 
alcohol: Brownish, dull sandy-mud. 

Remarks. — The types of Brada setosa Verrill from Massachusetts, 
B. pilosa Moore from Alaska, and Stylaroides plurihranchiata Moore 
from California were examined and are hereui referred to B. villosa. 
The excessive "hairiness" of B. pilosa appears to be a variable charac- 
ter; there may be varying numbers of papillae as well as all gradations 
in the amount of encrusting sand grains. Two large specimens washed 
ashore at Point Barrow were at first identified as B. rugosa, following 
St0p-Bowitz (1948); they agree in the large size (23-24 setigers, 28-30 
mm. long, 5-7 mm. wide), the almost complete absence of papulation, 
and the thick rugose surface; the outer surface, however, appears to be 
worn off. There are some remains of the papillated surface on the 
parapodial lobes. Thus B. rugosa (including T. arctica) is herein 
referred to B. villosa, where it has been referred earlier by Fauvel and 
others. 

New records. — Arctic Alaska: Point Barrow base, washed ashore 
September 22, 1949 (22 specimens), August 10, 1950 (2 specimens); off 
Point Barrow base, up to 4 mUes from shore, 27-29 fms., on various 
combinations of mud, gravel, stones, rocks (2 stations, 2 specimens), 
Bering Sea: 62°15' N., 167°48' W., Stoney, 1884; Albatross Sta. 3337, 
53°55' N., 163°26' W., 280 fms., 1890. West Coast North America: 
Washington Sound, Pettibone; Albatross Sta., northwest West Point, 
Elliot Bay, Seattle, 1914. East Coast North America: Off Nova 



292 PROCEEDINGS OF THE NATIONAL MUSEUM voi-.ios 

Scotia, Maine, Massachusetts, Rhode Island, 6-499 fms., U. S. Fish 
Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Barents Sea, Novaya 
Zemlya, Kara Sea. Also Iceland, Faroes, Norway to Spain, Mediter- 
ranean, Adriatic, south Arabian Coast; Hudson Bay to Rhode Island; 
Bering Sea to southern California; north Japan Sea to Japan; South 
Atlantic: South Orkney and South Shetland Islands. In low water 
to 853 fathoms. 

Brada inhabilis (Rathke, 1843) 

FiGUEE 33, h 

Siphonostoma inhahile Rathke, 1843, p. 218, pi. 11, fig. 13. 

Brada sublaevis Stimpson, 1854, p. 32.— Verrill, 1881, pp. 289, 304.— Webster 
and Benedict, 1887, p. 731. 

Brada granulata Malmgren, 1867, p. 85, pi. 12, fig. 71.— Th^el, 1879, p. 52.— 
not Murdoch, 1885, p. 155 ( = S. villosa). — Annenkova, 1937, p. 176; 1938, 
p. 186.— Hartman, 1944a, pp. 334, 341; 1948, p. 41.— Zatsepin, 1948, p. 
135, pi. 33, fig. 3.— Wesenberg-Lund, 1950a, p. 36; 1950b, p. 85. 

Brada granosa Treadwell, 1937, p. 32 (not B. granosa Stimpson, 1854). 

Brada inhabilis St0p-Bowitz, 1948a, p. 40, fig. 11; 1948b, p. 42, fig. 16.— Wesen- 
berg-Lund, 1951, p. 80. 

Description. — Length 30-60 mm., width 8-12 mm., segments 23-26. 
Body subfusiform, grub-shaped, flattened ventrally, strongly arched 
dorsally, usually curved ventrally (in alcohol) . Body surface covered 
with low globular papillae (to naked eye surface appears almost 
smooth, granular), usually with thin layer of sand grains, often with 
an almost uniform layer of larger sand grains on dorsum. Prostomium 
and buccal segment with two groups of tlu-ee to four filiform branchiae, 
pair of grooved palps, and buccal siphon around mouth. First seg- 
ment achaetous (may be few very, delicate capillary setae), second 
segment with two small neurosetae, rest of segments with neuropodia 
as slightly elevated lobes and about four (2-9) amber-colored nem-o- 
setae with curved tips. Without distinct notopodia ; notosetae poorly 
developed, one or two or absent, extremely capillary, easily overlooked. 
Nephridial papillae on ventral side between segments 4 and 5 (setigers 3 
and 4, since the first segment is usually achaetous). Color: In life: 
Orange yellow. In alcohol : Light gray to ashy. 

Remarks. — B. granulata Malmgren is herein referred to B. inhabilis, 
following St0p-Bowitz (1948a). B. sublaevis Stimpson should also be 
referred here and not to B. rugosa (Hansen) as indicated by St0p- 
Bowitz (1948a, p. 37). 

New records. — Arctic Alaska: Off Point Barrow base, up to 12.1 
miles from shore, 20-123.5 fms., on bottoms of rocks, stones, gravel, 
and mass of worm tubes (5 stations, 9 specimens). Northwest 



MARINE POLYCHAETE WORMS — PETTIBONE 293 

Greenland: 1 mile northwest Conical Rock, 25-60 fms., Bartlett, 
1938. Bering Sea: Albatross Sta. 3223, 54°26' N., 165°32' W., 
56 fms., 1890; Bering Strait, 13 fms., Dall, 1880. Alaska: Iliuliuk 
Harbor, Unalaska, 1871, and New Harbor, Unga Island, under stones, 
1872, Dall; Kodiak, W. J. Fisher. East Coast North America: 
Off Labrador, 95 fms.. Blue Dolphin 'Eixpedition, 1949; Bay of Fmidy, 
Maine, 48-90 fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Novaya Zemlya,Kara 
Sea. Also Iceland, Faroes, Norway to Danish waters; Labrador to 
Maine; Bering Sea to Gulf of Alaska; north Japan Sea. In low water 
to 609 fathoms. 

Family Scalibregmidae 

Body inflated anteriorly or short, fusiform. Integument roughened 
or tesseiated and divided into superficial rings. Prostomium small, 
bilobed, or entire and with frontal horns; two ciliated evaginable 
nuchal grooves on each side of prostomium (fig. 33, i). Buccal 
segment achaetous. Parapodia biramous, rami small, with or without 
dorsal and ventral cirri. Setae simple, capillary, some single, some 
forked. Branchiae limited to anterior few segments (may be absent). 
Pygidium with or without anal cirri. Proboscis eversible, unarmed, 
globular, smooth. 

Represented by a single species from Point Barrow. 

Genus Scalibregma Rathke, 1843 
Scalihregma inflatum Rathke, 1843 

FiGUEE 33, i-k 

Scalibregma inflatum Rathke, 1843, p. 184, pi. 9, figs. 15-21. — Moore, 1923, 
p. 217.— Fauvel, 1927, p. 123, fig. 44, a-f ; 1932, p. 187; 1934a, p. 50.— 
Augener, 1928, p. 754.— Monro, 1930, p. 163.— Annenkova, 1934, p. 322; 
1937, p. 176; 1938, p. 186.— Okuda, 1938b, p. 99.— Hartman, 1944a, pp. 
336, 341; 1948, p. 40.— St0p-Bowitz, 1946, p. 67, fig. 2; 1948b, p. 24, fig. 8.— 
Zatsepin, 1948, p. 136, pi. 33, fig. 10.— Wesenberg-Lund, 1950a, p. 36; 
1950b, p. 86; 1951, p. 81.— Berkeley and Berkeley, 1952, p. 58, figs. 119-121. 

Scalibregma brevicauda Verrill, 1873, pp. 416, 422, 605; 1881, p. 302, pi. 9, fig. 5. 

Scalibregma minutum Webster and Benedict, 1887, p. 727. 

Description. — ^Length up to 100 mm., width 13 mm., 45-60 segments 
(two specimens washed ashore after a storm were longer than any 
previously recorded). Body divided into two more or less distinct 
regions, anterior inflated portion (anterior 15-17 setigers) tapered 
anteriorly, without dorsal and ventral cirri, and narrower attenuated 
posterior region with dorsal and ventral cirri. A more or less devel- 



294 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

oped ventral groove along length of body, better developed in posterior 
region. Integument tesselated, with four annuli per segment. Pro- 
stomium subquadrangular, wider anteriorly, with frontal horns on 
anterolateral borders, with pair of evaginable nuchal organs lateral 
to prostomium (when inverted, appear as diagonal slits). Buccal 
segment achaetous, forming rugose ring around prostomium. Mouth 
ventral, at level of first setigerous lobes. Proboscis eversible, globular, 
smooth, unarmed. Branchiae four pairs, short, tufted, arborescent, 
inserted posterior to notopodia on setigers 2-5, increasing in size 
posteriorly. Noto- and neuropodial lobes of anterior segments small, 
projecting, subequal. Beginning on setigers 16-18, dorsal and ventral 
cirri develop gradually, becoming elongate-conical, flattened or 
inflated. Setae silky, iridescent, capillary (may be extra long in 
epitokous pelagic form). Lateral ciliated organs between rami. 
Pygidium with five^o seven, short to long, deciduous, filiform anal 
cirri on ventral side. Epitokous sexual forms may appear in surface 
waters, massed with sexual products, with extra long setae and 
usually with highly pigmented dorsal and ventral cirri. Color: 
In life : Orange yellow. In alcohol : Without color, orange yellow, or 
sulfur yellow with darker pigmented masses in the outer portions of 
the dorsal and ventral cirri. 

Remarks. — Scalibregma brevicauda Verrill is herein referred to S. 
injlatum. Although the types were not examined, specimens from 
the New England region identified by Verrill as S. brevicauda were 
observed. The types of S. minutum Webster and Benedict from 
Eastport, Maine, were examined and found to agree with S. injlatum. 

New records. — Arctic Alaska: Point Barrow base, washed ashore; 
off Point Barrow base, 22.5-78 fms,, up to 15 miles from shore, on 
bottoms of mud and various combinations of mud, sand, gravel, 
stones, rocks, with shells and worm tubes (9 stations, 17 specimens). 
West Coast North America: Washington and Puget Sounds, 25 
fms., mud, Pettibotie. East Coast North America: Off Labrador, 
17-35 fms.. Blue Dolphin Expeditions, 1950, 1951; off Nova Scotia, 
Maine, Massachusetts, Rhode Island, Connecticut, Long Island 
Sound, 5-95.5 fms., U. S. Fish Commission; Cuttyhunk, Massachu- 
setts, low water, in sand, Pettibone, 1950. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Jan Mayen, Spits- 
bergen, Franz Josef Land, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes, Norway to France, Adriatic, Gulf of Oman; Labrador to 
Long Island Sound; Bering Sea to central California; north Japan 
Sea to Japan. Southern latitudes: New Zealand, Magellan Straits, 
South Georgia, Prince Edward Islands, Kerguelen. In low water to 
1,333 fathoms; epitokes at surface. 



MARESTE POLYCHAETE WORMS — ^PETTIBONE 295 

Family Opheliidae 

Body relatively short, often grublike or lancet-shaped. Integument 
smooth or granulated; segmental grooves weak or obscured by super- 
ficial annuli. Prostomium small, conical, without appendages; pair 
of eversible nuchal organs (fig. 33, I, o). Proboscis eversible, saclike, 
unarmed. Parapodia biramous, lobes reduced or lacking; setae simple, 
capillary (fig. 33, m, q). Without dorsal cirri; rarely with ventral 
cirri. Branchiae paired, cirriform, inserted near parapodia, extending 
along most of body (may be lacking). Pygidium forming an anal 
cylinder, a flap, or knob, with or without cirri (fig. 33, n, p). 

Represented by two genera and two species from Point Barrow. 
Both genera have paired cirriform branchiae along most of length of 
body; without lateral eyes along body. 

Key to the genera of Opheliidae from Point Barrow 

1. Body lancet-shaped; ventral surface thick, solelike, formed by 2 thick longi- 
tudinal muscular columns, with midventral and lateral grooves along length 
of body. Parapodial lobes small, along lateral grooves (fig. 33, m). 

Ammotrypane (p. 295) 
Body cylindrical, grub-shaped, without midventral or lateral grooves. Para- 
podial lobes absent, the simple, capillary noto- and neurosetae emerging 
directly from body wall (fig. 33, q) Travisia (p. 296) 

G^nus Ammotrypane Ratlike, 1843 
Ammotrypane breviata Ehlers, 1913 

Figure 33, l-n 

Ammotrypane breviata Ehlers, 1913, p. 523, pi. 39, figs. 1-7. — Monro, 1930, p. 

165; 1936, p. 165. 
Ophelina groenlandica St0p-Bowitz, 1948b, p. 20, fig. 6. 

Description. — Length 6-7 mm., width 0.3 mm., setigerous segments 
26 or 27 (up to 34 mm. long, 2 mm. wide, 26-28 segments — Monro, 
1930, 1936). Body rigid, tapered at both ends, smooth, without 
superficial segmental divisions, Prostomium conical, the anterior tip 
constricted to form a small globular palpode; prostomial tip may be 
partly telescoped into rest of prostomium; pair of eversible nuchal 
organs at level of parapodia. Parapodia with two groups of fine 
capillary setae, the dorsal group slightly longer than the ventral. 
Branchiae begin on setiger 2, absent on last three to five setigers; 
branchiae slender, cirriform, subequal. Pygidium forming a short, 
somewhat contractile anal cylinder, closed on aU sides, somewhat 
lobulate or papillate on open posterior end. Eversible proboscis 
saclike, smooth. Color: In alcohol: Without color or may be darkly 
pigmented laterally along posterior three setigerous segments and 
proximal part of anal tube. 



296 PROCEEDINGS OE THE NATIONAL MUSEUM vol. 103 

Remarks. — The two small specimens from Point Barrow (6-7 mm. 
long) follow closely the description of St0p-Bowitz for Ophelina groen- 
landica, including size. They seem to agree with the description by 
Ehlers of A. breviata from the Antarctic, although the Antarctic 
specimens attain a much larger size (29 mm. long). 

New records. — Arctic Alaska: Off Point Barrow base, up to 7.5 
miles from shore, 29-36 fms., on bottoms of gravel, stones, large 
perforated rocks (2 stations, 2 specimens). 

Distribution. — ^Arctic Alaska, East Greenland. Southern latitudes: 
South Orkney and Falkland Islands, Kaiser- WUhelm II Land. In 
10-214 fathoms. 

Genus Travisia Johnston, 1840 
Travisia camea Verrill, 1873 

Figure 33, o-q 

Travisia cornea Verrill, 1873, pp. 431, 434, 508, 604; 1881, p. 302, pi. 8, fig. 1.— 
Sumner, Osburn, and Cole, 1913, p. 629.— Hartman, 1944a, pp. 336, 341, pi. 
19, fig. 6 (as probably Ophelia limacina). 

Travisia forbesi Murdoch, 1885, p. 154 (not T. forbesii Johnston, 1840). 

Description. — Length up to 59 mm., width 8 mm., segments 25-29 
(specimens washed ashore at Point Barrow much larger than any 
previously recorded). Body rounded, cylindrical, grub-shaped, 
tapered toward each end. Segments of posterior third may be some- 
what imbricated or telescoped; triannulate in middle region. Integu- 
ment with granulations or beading rather uniform (not larger on 
posterior part of segment as in T. pupa Moore). Prostomium small, 
conical, acutely to bluntly pointed, without eyes. Branchiae begin on 
setiger 2 (rarely on setiger 3), continuing posteriorly, absent on last 
four or five segments, cirriform, inserted posterior to notosetae. 
Without parapodial lobes, the small, slender, capillary noto- and neuro- 
setae emerging directly from body wall. In posterior third of body, 
area in region of setae may be somewhat inflated but without enlarged 
lateral lobes extending posteriorly (as in T. forbesii Johnston and 



Figure 33. — Cirratulidae: a, Cirratulus cirratus, dorsal view anterior end; h, same, para- 
podium from anterior region; c, same, acicular crotchet; d, Chaetozone setosa, dorsal 
view anterior end. Flabelligeridae: e, Flabelligera affinis, parapodium; /, same, hooked 
pseudoconipound neuroseta; g, same, pedunculate papillae — (1) from tip of notopodium, 
(2) on body; k, Brada inhahilis, ventral view anterior end. Scalibregmidae: i, Scalihregma 
inflatum, dorsal view anterior end; /, same, parapodium from middle region; k, same, 
parapodium from posterior region. Opheliidae: /, Ammotrypane breviata, ventral view 
anterior end, proboscis extended; w, same, lateral view middle part of body; n, same, 
ventral view posterior end; o, Travisia camea, dorsal view anterior end; p, same, dorsal 
view posterior end; q, same, parapodium. Capitellidae: r, Capitella capitata, dorsal view 
anterior end; /, same, dorsal view posterior part thoracic region; t, same, parapodium 
from abdominal region; u, same, hooded crotchet. (For explanation of symbols, see 
p. 210.) 



MARINE POLYCHAETE WORMS — PETTIBONE 



297 




P Q 

Figure 33. — For explanation see facing page. 



298 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

T. brevis Moore). With lateral sensory pits between rami appearing 
as small oval slits. Nephridial pores ventral to nem-opodia on setigers 
3-14. Pygidium small, cylindrical, longitudinally fmrowed. Color: 
In life: Flesh. In alcohol: Whitish. 

New records. — Arctic Alaska: Point Barrow base, washed ashore, 
October 16, 17, 1949; August 10, 1950 (12 specimens). East Coast 
North America: Off Massachusetts, Long Island Sound, 3-19 fms., 
soft mud, U. S, Fish Commission; Woods Hole region, Massachusetts, 
Pettibone, 1950. 

Distribution. — Arctic Alaska. Massachusetts to Long Island Sound. 
In 3-19 fathoms. 

Family Capitellidae 

Body cylindrical, wealdy divided into 2 regions: Anterior thoracic 
region somewhat inflated, often reticulated, parapodial lobes rudi- 
mentary; posterior abdominal region longer and thinner, with rows 
of crotchets on somewhat inflated tori. Prostomium conical, more 
or less retractile, without appendages, with or without eyes, with 
pair of eversible nuchal organs. Buccal segment achaetous (may be 
fused with prostomium as in Capitella). Parapodia biramous, re- 
duced to bundles of capillary setae or to dorsal and ventral tori 
bearing row of crotchets which have long manubrium and recurved 
hooded rostrum (fig. 33, r-^). Without dorsal, ventral, or anal cirri. 
With or without branchiae. Proboscis eversible, globular, unarmed, 
papillate. Constructs tubes. 

Represented by a single species from Point Barrow. 

Genus Capitella Blainville, 1828 
Capitella capitata (Fabricius, 1780) 

Figure 33, r-u 

Lumbricus capitatus Fabricius, 1780, p. 279. 

Capitella capitata Webster and Benedict, 1884, p. 730; 1887, p. 744. — Ehlers, 
1913, p. 543.— Eliason, 1920, p. 63.— Fauvel, 1927, p. 154, fig. 55, a-h.— 
Augener, 1928, p. 749.— Monro, 1930, p. 163.— Anneukova, 1934, p. 322; 
1937, p. 179; 1938, p. 189.— Hartman, 1942a, p. 69; 1944a, pp. 334, 341, 
pi. 34, fig. 3; 1945, p. 37; 1947, p. 404, pi. 43, figs. 1, 2; 1948, p. 41; 1951, 
p. 101.— Berkeley and Berkeley, 1943, p. 129; 1952, p. 100, figs. 206-208.— 
Gorbunov, 1946, p. 39.— Thorson, 1946, p. 110, fig. 60.— Zatsepin, 1948, 
p. 138, pi. 33, fig. 14. — Hartman and Reish, 1950, p. 40. — Wesenberg-Lund, 
1950a, p. 39; 1950b, p. 91; 1951, p. 84. 

Capitella capitata var. antarctica Monro, 1930, p. 164. 

Description. — Length 23 mm., width 1 mm. (up to 120 mm. — 
Augener, 1928). Body attenuated slightly anteriorly, much more 
so posteriorly, fragile. Prostomium and achaetous buccal segment 
fused, forming wide obtuse cone, somewhat flattened dorsoventrally, 



MARINE POLYCHAETE WORMS — PETTIBONE 299 

usually concave dorsally/witli''pair^of small ventral eyes and pair of 
crescent-shaped nuchal slits behind eyes (best seen from lateral view) , 
Thoracic region smooth, without reticulated integument, consisting 
of first 9 setigerous segments, each of first 7 with four bundles of 
capUlary setae which on setiger 7 may be only capillaries, only crotch- 
ets, or both (in young specimens, crotchets may be present from 
setiger 4) . Setigers 8 and 9 modified, having neuropodia with row of 
hooded crotchets, and with large, recurved, amber-colored acicular 
genital hooks located middorsally between these two segments. 
Abdominal segments with slightly raised dorsal and ventral para- 
podial tori each bearing a row of hooded crotchets. Without spe- 
cialized branchiae or lateral organs. Pygidium a short ring, may be 
inflated. Color: In alcohol: Without color or reddish brown. 
Females with large yolky eggs (Point Barrow, August 9, 30, 1949; 
September 1, 6, 1949.) 

Remarks. — According to Fauvel and others, in C. capitata the genital 
hooks on setigers 8 and 9 are present only in the males, the females 
having only crotchets. On all the specimens from Point Barrow, 
genital hooks are present, including some females containing large 
yolky eggs. They agree in all other respects with C. capitata. Having 
genital hooks in both males and females, they would fall under the 
very closely related Capitellides Mesnil. They differ from C. giardi 
Mesnil in the larger number of genital hooks. They are herein re- 
ferred to the cosmopolitan C. capitata with the tentative idea that the 
relative development of genital hooks in the female may prove to be 
a variable character. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, up to 7.5 miles from 
shore, 23.3-70 fms., on bottoms of stones and various combinations 
of gravel, stones, rocks, large perforated rocks (8 stations, 26 speci- 
mens). Bering Sea: St. Paul Island, Pribilofs, in holdfasts with 
Fabricia sabella and lumbrinerids, W. L. Hahn, 1911. West Coast 
North America: Corona del Mar, California, from branchial cavity 
of crab, Loxorhynchus grandis, MacGinitie, 1948. East Coast North 
America: Off Labrador, 6 fms., mud-sand. Blue Dolphin Expedition, 
1949; off Massachusetts, 15 fms., U. S. Fish Commission; Woods Hole 
region, Massachusetts, Pettibone, 1951. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Spitsbergen, Novaya 
Zemlya, Kara Sea. Also Iceland, Faroes, Norway to France, Madeira, 
Mediterranean, Black Sea; Hudson Bay to North Carolina, Texas; 
Bering Sea to southern California; north Japan Sea. Southern lati- 
tudes : Magellan Straits, South Georgia, Bouvet Island, South Africa, 
Kerguelen. In low water to 500 fathoms. 

261112—54 7 



300 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Family Arenicolidae 

Body elongate, cylindrical, integument reticulated and divided into 
secondary annuli. Body divided into two or three distinct regions: 
(1) anterior, abranchial; (2) abdominal, branchial, each segment with 
pair of dorsally-borne branched gills; (3) caudal, achaetous and 
abranchial; this region may be lacking. Prostomium small, simple 
or trilobed, without appendages; with nuchal groove (fig. 34, a). 
Parapodia biramous; notopodia conical, setigerous, with spiny, capU- 
lary setae; neuropodia with sigmoid crotchets in semicircular tori. 
Without dorsal, ventral, and anal cirri. Anus terminal. Proboscis 
eversible, globular, papillate, unarmed. Constructs tubes or burrows. 

Represented by a single species from Point Barrow. 

Genus Arenicola Lamark, 1801 
Arenicola glacialis Murdoch, 1884 

Figure 34, a 

Arenicola glacialis Murdoch, 1884, p. 522; 1885, p. 155. — Ashworth, 1910, p. 24, 
figs. 10-14; 1912, p. Ill, figs. 46, 47, pi. 6; 1924, p. 3, fig. 1. 

Description. — Length up to 205 mm., width 11 mm., 17 setigerous 
segments. Body cylindrical, tapering slightly anteriorly and gradu- 
ally posteriorly. Prostomium small, may be pretty well retracted in 
nuchal groove; median lobe small, lateral lobes larger, curved (fig. 
34, a). Body divided into three regions: (1) anterior region of six 
abranchial setigers, notopodia conical with spiny capillary setae, 
neuropodia low tori with few sigmoid crotchets; (2) branchial region 
of 11 setigers each divided into five superficial annuli, with short, 
bushy, arborescent branchiae at base of conical notopodia, with gill 
axes short, branching one or two times or simple; neuropodia raised 
tori with long row of crotchets; on more posterior segments, tori 
approach midventral line; (3) caudal achaetous and abranchial region 
with papillae feebly developed, without processes (as in A. cristata). 
Nephridial pores six pairs, on setigers 4-9, posterior to upper part of 
neuropodial tori. Pair of statocysts on peristomium, open to exterior. 



Figure 34. — Arenicolidae: a, Arenicola glacialis, prostomium. Maldanidae: b, Praxillella 
praeUrmissa, ventral view anterior end; c, same, frontal view cephalic plate; d, same, 
pygidial funnel; g, same, posterior end; /, same, crotchet from neuropodium; g, Maldane 
sarsi, lateral view anterior end; h, same, lateral view posterior end; i, Nicomache lum- 
bricalis, lateral view anterior end; /, same, lateral view posterior end; k, Nicomache 
per sonata, lateral view posterior end; /, Petaloproctus tenuis, lateral view anterior 
end; m, same, lateral view posterior end. Sabellariidae: n, Idanthyrsus armatus, dorsal 
view anterior end; o, same, lateral view anterior end; p, same, palea from inner row; 
q, same, palea from outer row. (For explanation of symbols, see p. 210.) 



MARINE POLYCHAETE WORMS — PETTIBONE 301 

■ set 19, 




Figure 34. — For explanation see facing page. 



302 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

with numerous sand grains as statoliths. Color : In life : Deep orange 
to brownish red. In alcohol: Bluish to grayish black or brown. 

New records. — Arctic Alaska: Point Barrow base, washed ashore 
August 21, 24, 28, September 6, 12, 19, 22, October 16, 1949 (54 
specimens); off Point Barrow base, along shore, 1.7 fms., on bottom 
of gravel with mud (2 stations, 3 specimens) . 

Distribution. — Alaskan and Canadian Arctic. In 1.7-3 fathoms. 

Family Maldanidae 

Body cylindrical, truncated at both ends, without differentiated 
regions, segments usually few, some much longer than wide. Pro- 
stomium a small cephalic crest fused with achaetous buccal segment, 
forming ovoid head, without appendages, with or without oblique 
limbate brim or plate (fig. 34, b, i, I) ; pair of nuchal slits on either side 
cephalic crest. In anterior region with numerous glandular cells, 
diffused or grouped in clear bands alternating with pigmented rings. 
Parapodia biramous; notopodia with capillary setae; neuropodia with 
projecting tori bearing rows of crotchets (fig. 34,/) with dentate ros- 
trum and rostral hairs; the crotchets may be replaced by few stout 
acicular spines on first few setigers. Without dorsal or ventral cirri; 
usually without branchiae. Pygidium a limbate plate, spatulate or 
funneUike (fig. 34, h, j, m). Proboscis eversible, unarmed, globular. 
Live in tubes in sand or mud; tube cylindrical, membranous, covered 
with thin to thick layer of mud, sand, gravel, more or less resistant. 

Represented by four genera and five species from Point Barrow. All 
the genera are without specialized collarettes, without special glandular 
ventral shield on setiger 8, with neuropodial crotchets in single rows. 

Key to the genera of Maldanidae from Point Barrow 

1. Head with limbate plate (fig. 34, h, g) 2 

Head without limbate plate (fig. 34, i, I). With 1-2 stout acicular setae in 
neuropodia of first 3 setigers. Pygidium funnel-shaped or forming concave 
plate, with or without cirri; anus opening on surface of plate 3 

2. Head with low cephalic crest, bordered by oblique limbate plate, entire or 

notched laterally (fig. 34, h, c) . With ventral crotchets beginning on setiger 1. 
Pygidium funnel-shaped, bordered with cirri, with anus in center of somewhat 

protruding anal cone within the funnel (fig. 34, d) Praxillella (p. 303) 

Head with middorsal convex cephalic crest or keel, bordered by limbate plate, 
divided into 3 parts by 2 deep lateral grooves (fig. 34, g). Without neuro- 
setae on setiger 1 ; crotchets begin on setiger 2. Pygidium a slightly oblique 
oval plate, bordered by rim which is incised laterally, smooth or denticled; 
anus dorsal to pygidial disc (fig. 34, h) Maldane (p. 303) 

3. Pygidium sHghtly obhque, funnel-shaped, bordered with cirri (fig. 34, j, k). 

Nicomache (p. 304) 

Pygidium an asj'^mmetrical, foliaceous, concave plate, reduced dorsally, wider 

ventrally, with border smooth or crenulate (fig. 34, m) .Petaloproctus (p. 306) 



MARINE POLYCHAETE WORMS — PETTIBONE 303 

Genus Praxillella Verrill, 1881 
Praxillella praetertnissa (Malmgren, 1865) 

Figure 34, b-f 

Praxilla praetermissa Malmgren, 1865, p. 191; 1867, p. 100, pi. 11, fig. 62. 
Praxillella praetermissa Verrill, 1881, pp. 298, 305, 309, 312.— Webster and 

Benedict, 1887, p. 746.— Arwidsson, 1907, p. 192, figs.— Eliason, 1920, p. 

66.— Augener, 1928, p. 762.— Annenkova, 1937, p. 182; 1938, p. 194.— 

Zatsepin, 1948, p. 142, pi. 35, fig. 1.— Wesenberg-Lund, 1948, p. 41, figs. 

20, 21; 1950a, p. 43; 1950b, p. 98; 1951, p. 93. 
Clymene {Praxillella) praetermissa Fauvel, 1927, p. 179, fig. 62, a-e. 

Description. — Length 45 mm., width 1 mm, (up to 120 mm. long, 
2.5 mm. wide — Malmgren, 1865), 19 setigerous segments, 4-5 achaetous 
segments with rudimentary parapodia. Head with oblique limbate 
plate, with border entire, slightly or deeply incised laterally, slightly 
incised posteriorly, terminating anteroventraUy in obtuse cone (not 
produced into digitiform process); median cephalic crest long; nuchal 
grooves one-half to two-thirds its length. Neuropodia of anterior 
three setigers with 2-10 slightly modified crotchets. Pygidial funnel 
with flat floor, with projecting anal cone with larger valvule on ventral 
side of cone; border of funnel with 20-38 subequal papillae except 
for a longer ventral one. Proboscis papillate basally. Color: In 
life: Yellow with wide, red and white bands on some anterior segments. 
In alcohol: Brownish to gray, with whitish bands anterior to setae 
on setigers 4-8. Tube : Of clay, sand, small bits of rock; may be quite 
resistant. 

New records. — Arctic Alaska: Off Point Barrow base, up to 7.5 
miles from shore, 20-36 fms., on various combinations of mud, sand, 
gravel, stones, rocks, large perforated rocks, shells (7 stations, 10 
specimens). East Coast North America: Off Labrador, 5-8 fms., 
Blv£ Dolphin Expedition, 1949; off Cape Cod, Massachusetts, 14-19 
fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Alaskan Arctic, 
Davis Strait, Greenland, Spitsbergen, White Sea, Novaya Zemlya, 
Kara Sea. Also Iceland, Faroes to English Channel, Mediterranean; 
Labrador to Massachusetts; north Japan Sea. In 7-1,111 fathoms. 

Genus Maldane Grube, 1860 
Maldane sarsi Malmgren, 1865 

Figure 34, g, h 

Maldane sarsi Malmgren, 1865, p. 188; 1867, p. 99, pi. 10, fig. 57. — Arwidsson, 
1907, p. 251, figs.— Eliason, 1920, p. 66.— Fauvel, 1927, p. 197, fig. 69, a-i; 
1932, p. 202.— Augener, 1928, p. 759.— Annenkova, 1937, p. 179; 1938, p. 
192.— Okuda, 1939, p. 239.— Berkeley and Berkeley, 1942, p. 199.— Hartman, 



304 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

1944a, pp. 335, 342, pi. 33, figs. 3, 4; 1951, p. 106.— Zatsepin, 1948, p. 142, 

pi. 33, fig. 3.— Wesenberg-Lund, 1948, p. 48, figs. 24, 25; 1950a, p. 44; 1950b, 

p. 100; 1951, p. 94. — Hartman and Reish, 1950, p. 37. 
Maldane sarsi var, antardica Arwidsson, 1911a, p. 32, figs. — Munro, 1930, p. 

169; 1936, p. 168. 
Maldane sarsi var. Moore, 1923, p. 237. 
Maldane sarsi var. tropica Monro, 1937, p. 307, fig. 24. 

Description. — Length up to 110 mm., width 3.9 mm., setigerous 
segments 19, two antanal achaetous segments. Head with long, 
convex keel, ending in blunt process above mouth, bordered by limbate 
plate, with border smooth, usually incised laterally, rather high pos- 
teriorly; nuchal organs short, divergent; without ocelli. Anterior 
segments biannulate. With notosetae only on first setiger; neuro- 
podial crotchets begin on setiger 2. Two antanal segments short, 
with achaetous tori slightly marked. Pygidium an oblique flat, oval 
plate, with border usually slightly incised laterally, smooth or slightly 
crenulate ventrally. Anus a folded cushion dorsal to and close to 
pygidial rim. Tube: Free, circular, thin to thick walled, of fine 
agglomerated mud and sand grains. 

New Records. — Arctic Alaska: Off Point Barrow base, 3 miles 
from shore, 20.3 fms., on bottom of mud, gravel, stones, shells (1 
station, 1 specimen), Bering Sea: Albatross Sta. 3603, 55°23' N., 
170°31' W., 1,771 fms., 1895. Canadian Arctic: Northwest shore 
Daniels Island, mouth Newell Sound, Frobisher Bay, Baffin Land, 
10-30 fms., Bartlett. East Coast North America: Off Labrador, 
30 fms.. Blue Dolphin Expedition, 1950; off Nova Scotia, Maine, 
Massachusetts, Rhode Island, 14-487 fms., U. S. Fish Commission., 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan 
and Canadian Arctic, Davis Strait, Greenland, Spitsbergen, Novaya 
Zemlya, Kara Sea. Also Iceland, Norway to France; Labrador to 
Rhode Island, Louisiana, Mississippi, eastern Texas; Bering Sea to 
southern California; north Japan Sea to Japan; South Arabian coast 
(var. tropica); Bay of Bengal. Southern latitudes: South Georgia, 
South Shetlands, Palmer Archipelago (var. antarctica) ; south Austraha. 
In low water to 1,771 fathoms. 

Genus Nicomache Malmgren, 1865 

Both species have buccal segment fused with prostomium forming 
hood-shaped head, irregularly rounded, with thickened anteroventral 
border, forming projecting upper lip (fig. 34, i). First three setigerous 
segments each with one or two stout, acicular, spinelike nem"0setae. 
Some of notopodia with long, hairlike sinuous dorsal setae. Pygidium 
with anus terminal in center of short, slightly oblique funnel, with 
papillae on border of funnel (fig. 34, j, k) . 



MARINE POLYCHAETE WORMS — PETTIBONE 305 

Key to the species of Nicomache from Point Barrow 

1. Single achaetous antanal segment (fig. 34, k) N. personata 

Two achaetous antanal segments (fig. 34, j) N. lumbricalis 

Nicomache lumbricalis (Fabricius, 1780) 

Figure 34, i, j 

Sabella lumbricalis Fabricius, 1780, p. 374. 

Nicomache lumbricalis Malmgren, 1867, p. 99, pi. 10, fig. 60. — Webster and 
Benedict, 1884, p. 731; 1887, p. 745.— Arwidsson, 1907, p. 86, pi. 8, figs. 
244, 245.— Fauvel, 1927, p. 190, fig. 66, a-i.— Augener, 1928, p. 764.— Monro, 
1930, p. 173.— Annenkova, 1934, p. 322; 1937, p. 181; 1938, p. 193.— Hartman, 
1944a, pp. 335, 342; 1948, p. 42.— Zatsepin, 1948, p. 144, pi. 26, fig. 1.— 
Wesenberg-Lund, 1948, p. 23, figs. 10, 11; 1950a, p. 41; 1950b, p. 95; 1951, 
p. 90.— Berkeley and Berkeley, 1952, p. 54, figs. Ill, 112. 

Nicomache carinaia Moore, 1906b, p. 242, pi. 11, figs. 36-39, pi. 12, figs. 43, 44; 
1923, p. 227. 

Nicomache lumbricalis var. borealis Arwidsson, 1907, p. 94, figs.; 1922, p. 6. — 
Eliason, 1920, p. 65. 

Description. — Length 40-60 mm., width 2.5-3 mm., setigerous 
segments 21-22, two short achaetous antanal segments (Point Barrow 
and Labrador specimens with 22 setigers; type of N. carinaia with 21 ; 
up to 160 mm. long, 5 mm. wide — Fauvel, 1927). Prostomium with- 
out ocelli; nuchal organs S-shaped. Pygidial funnel with 14-30 
subequal, pointed, triangular papillae. Color: In alcohol: Reddish 
brown on sides and dorsal surface of head and first three setigers. 
Tube: Sandy, with small pieces of rock, with smooth membranous 
lining (Point Barrow), or thick, sandy, and coiled (Labrador; small 
ones on terebeUid tubes) . 

Remarks. — They may share their tubes with the commensal poly- 
noid Enipo gracilis Verrill (=E. cirrata TreadweU) reported by Berke- 
ley and Berkeley from Alaska; also from off Nova Scotia and Mas- 
sachusetts (in USNM). 

New records. — Arctic Alaska: Off Point Barrov/, up to 12.1 miles 
from shore, 24.7-123.5 fms., on bottoms of mud, worm tubes, stones, 
and various combinations of rocks, stones, gravel (6 stations, 18 
specimens). Bering Sea: Cape Prince of Wales, 23 fms., Dall, 1874. 
Albatross Sta. 3311, 53° 59' N., 166° 29' W., 85 fms., and Sta. 3313, 
54° or N., 166° 27' W., 68 fms., 1890. Alaska: Albatross Sta. 2848, 
Unalaska to Cook Island, 1888; Eastern Harbor, Sitka, 15 fms., Dall, 
1874. Kamchatka: Albatross Sta. 3644, 51° 09' N., 157° 48' W., 96 
fms., 1896. East Coast North America: Off Labrador, 30-125 
fms.. Blue Dolphin Expeditions, 1949, 1950, 1951; off Nova Scotia, 
Bay of Fundy, Maine, New Hampshire, Massachusetts, 26-150 fms., 
U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian and 



306 PROCEEDINGS OF THE NATIONAL RIUSEUM vol. 103 

Alaskan Arctic, Greenland, Jan Mayen, Spitsbergen, White Sea, 
Novaya Zemlya, Kara Sea, Also Bering Sea to Lower California; 
north Japan Sea; Iceland, Norway to English Channel, Santander; 
Labrador to Massachusetts; South Africa. In low water to 1,400 
fathoms. 

Nicomache personata Johnson, 1901 

Figure 34, k 

Nicomache personata Johnson, 1901, p. 419, pi. 13, figs. 134-139. — Arwidsson, 

1922, p. 7, pi. 1, figs. 5, 6.— Hartman, 1948, p. 41, fig. 11, d-g.— Berkeley 

and Berkeley, 1952, p. 54, figs. 109, 110. 
Nicomache minor Arwidsson, 1907, p. 100, pi. 2, figs. 68-73; pi. 8, figs. 252-256. — 

Augener, 1928, p. 765.— Annenkova, 1937, p. 181; 1938, p. 193. 
Nicomache maculata Arwidsson, 1911b, p. 209, pi. 18, figs. 13-19; pi. 19, figs. 

27-30.— Fauvel, 1927, p. 191, fig. 66, k-r. 

Description. — Up to 110 mm. long, 2 mm. wide, 22 or 23 setigerous 
segments, single achaetous antanal segment (Point Barrow specimen 
with 23 setigers, Puget Sound specimen with 22). Prostomium with 
numerous ocelli (may be absent) ; nuchal clefts short, arched. Pygidial 
funnel with 16-26 papillae which are short, subequal, triangular, or 
very unequal, some being longer, some very short, few may be bifid 
or trifid. Color: In alcohol: Without color, or brownish, especially 
on anterior end. Tube : Thick walls of coarse sand particles cemented 
together. 

New records. — Arctic Alaska: Off Point Barrow base, up to 8 
miles from shore, 21.7-75.5 fms., on various combinations of pebbles, 
rocks, gravel, stones (3 stations, 5 specimens). Bering Sea: Alba- 
tross Sta. 3289, 56° 44' N., 159° 16' W., 16 fms., 1890. West Coast 
North America : Security Bay, Alaska, Jones; Washington and Puget 
Sounds, low water to 61 fms., Pettiboiie. 

Distribution. — Arctic Alaska, Spitsbergen; Bering Sea to Washing- 
ton; north Japan Sea; Scandinavia, Ireland, North Sea to English 
Channel. In low water to 75.5 fathoms. 

Genus Petaloproctus Quatrefages, 1865 

Petaloproctus tenuis (Theel, 1879) 

Figure 34, I, m 

Maldane tenuis Th^el, 1879, p. 57, pi. 4, figs. 52-54. 

Maldane filifera Verrill, 1879, p. 179. 

Petaloproctus tenuis Arwidsson, 1907, p. 114, pi. 6, fig. 190a. — Annenkova, 1937, 

p. 181; 1938, p. 193.— Zatsepin, 1948, p. 145, pi. 34, fig. 5.— Wesenberg-Lund, 

1948, p. 31.— Berkeley and Berkeley, 1952, p. 55, fig. 113. 
Petaloproctus tenuis var. borealis Arwidsson, 1907, p. 118, pi. 3, figs. 85-90; pi. 8, 

figs. 268-272.— Eliason, 1920, p. 66.— Hartman, 1948, p. 42.— Berkeley and 

Berkeley, 1952, p. 56, figs. 114-116. 
Petaloproctus filifer Hartman, 1942b, p. 131, fig. 11, a-b. 



MARINE POLYCHAETE WORMS — PETTIBONE 307 

Description. — Length 18 mm., width 0.5 mm., 20-21 setigerous 
segments, smgle achaetous antanal segment (may appear as two or 
three on longer ventral side). Prostomium rounded, with convex 
median ridge and anteroventraUy extended brim. First three setigers 
with one or two acicular nem"0setae; with some long, threadlike, sinu- 
ous notosetae covered with spinules. Pygidium with obUque, slightly 
concave, disclike, asymmetrical plate (absent dorsally), wide on ven- 
tral side, with rim smooth or slightly crenulate; anal opening terminal. 

Remarks. — The Point Barrow specimen agrees with the stem form 
of P. tenuis, with 20 setigerous segments and the rim of the pygidial 
disc slightly crenulate (in var. horealis and Jilifer there are 21 setigers 
and the rim of the pygidial disc is smooth) , 

Nev) records. — Arctic Alaska: Off Point Barrow base, 16 miles 
from shore, 78.2 fms., on bottom of worm tubes, few rocks (1 station, 
1 specimen). Canadian Arctic: Ducketts Cove, Hurd Channel, 
Melville Peninsula, 1-14 fms., in sandy tubes on empty tubes of 
Pectinaria, Bartlett, 1933. Greenland: Small bay. Camp No. 2, 
Nugsuak Peninsula, 74°7' N., on empty tubes of Pectinaria, J. C. 
Martin, 1897. 

Distribution. — Scattered records in the Arctic: Alaskan and Cana- 
dian Arctic, East Greenland, Spitsbergen, Novaya Zemlya. Also off 
Cape Cod, Massachusetts; southeastern Alaska to British Columbia; 
north Japan Sea. In 1-833 fathoms. 

Family Sabellariidae 

Body divided into three regions: (1) anterior thoracic region includ- 
ing two anterior thoracic segments with parapodia reduced to neuro- 
podia with capillary setae and two large, more or less fused, heavy 
columns, the opercular peduncles, directed anteriorly, bearing very 
modified setae arranged in one to three concentric rows and forming an 
operculum for closing opening of tube; three or four para thoracic 
biramous segments with capillary neurosetae and paddlelike or 
styliform notosetae and dorsal falciform branchiae (fig. 34, n-q) ; (2) 
abdominal region with notopodia in form of broadly flattened pinnules 
bearing pectinate uncini, and neuropodia with capillary setae; with 
dorsal, simple, ligulate or falciform branchiae; (3) caudal region, 
achaetous, apodous, cylindrical, without appearance of segmentation. 
Prostomium indistinct, between opercular peduncles, with pair of 
palps and usually a median tentacle. Mouth unarmed, usually sur- 
rounded with numerous filiform tentacles. Tube of sand or fine 
gravel, concreted, resistant, may be solitary or colonial, constructing 
sandy reefs. 

Represented by a single species from Point Barrow. 



308 PROCEEDINGS OF THE NATIONAL MUSEUM voi^ m 

Genus Idanthyrsus Kinberg, 1867 
Idanthyrsus armatus Eauberg, 1867 

Figure 34, n-q 

Idanthyrsus armatus Kinberg, 1867, p. 350. — Johansson, 1926, p. 9; 1927, p. 90. — 
Monro, 1930, p. 117, fig. 73; 1936, p. 172.— Annenkova, 1937, p. 184; 1938, 
p. 196.— Okuda, 1938a, p. 242, figs. 4, 5.— Hartman, 1944c, p. 336, pi. 31, 
fig. 36.— Berkeley and Berkeley, 1952, p. 107, figs. 220-222. 

Idanthyrsus ornamentatus Chamberlin, 1919, p. 262, pi. 3, figs. 2-5. — Hartman, 
1944c, p. 337, pi. 31, fig. 34; 1948, p. 43.— Hartman and Reish, 1950, p. 41. 

Description. — Length up to 60 mm., width 5 mm., caudal lobe 11 
mm. Body widest anteriorly, tapering gradually posteriorly, with 
achaetous caudal region sharply set off from body. Opercular pe- 
duncles elongated, semicyUndrical, completely separated and diver- 
gent anteriorly, obliquely truncated distally, with two rows modified 
golden setae or paleae; inner row paleae rather stout, smooth, gently 
curved, tapering gradually to acute tips, 7-16 per column; outer row 
paleae more slender, straight or slightly curved, with coarse lateral 
spines, 16-36 per column; row of 9-15 distal papillae per column, at 
base of outer paleae. One or two pairs heavy, deep amber-colored, 
strongly-bent nuchal hooks on dorsal side opercular peduncles. 
Numerous filiform oral tentacles on ventral side opercular peduncles. 
Pair of grooved slender palps and median tentacle between bases of 
peduncles. First thoracic segment short, with ventral group capiUary 
setae lateral to conical papilla. Second segment with lateral group 
capillary setae, three papillae and dorsal branchiae. Three para- 
thoracic segments with rectangular palletlike pinnules with flattened, 
paddlelike ilotosetae; neurosetae bipinnate capillary setae. Abdom- 
inal segments with wide, flattened dorsal pinnules bearing pectinate 
uncini; capillary barbed neurosetae. Caudal lobe smooth, achaetous. 
Paired dorsal branchiae cirriform, with transverse ridges on medial 
side, begin on setiger 2, absent on last few abdominal segments. 
Color: In life: Reddish brown to dark violet anteriorly, branchiae 
red, paleae golden yellow. In alcohol: Colorless, with purple 
patches anteriorly. Tube: Thick, of coarse sand grains neatly ce- 
mented together, very resistant, one side flattened, attached to rocks, 
shells, crustaceans; upper surface convex. 

Remarks. — /. ornamentatus Chamberlin is herein referred to /. 
armatus, following Okuda (1938). According to observations by G. 
E. MacGinitie it is a hardy species, a specimen living in a pail in the 
laboratory for three days, on a rock covered with a mass of barnacles, 
etc. Male specimen, collected from 70 fathoms, spawned in the 
laboratory August 9, 1949. 



MARINE POLYCHAETE WORMS — PETTIBONE 309 

New records. — Arctic Alaska: Off Point Barrow base, up to 7 
miles from shore, 4.3-70 fms., on bottoms of mud, and gravel, stones 
(3 stations, 6 specimens) . Bering Sea : Bering Island, N. Grebiatzky, 
1884. West Coast North America: Albatross Sta. 2842, 54°15' N., 
166°3' W., 72 fms.; Washington and Puget Sounds, 10-110 fms., 
Pettibone. 

Distribution. — Arctic Alaska, Bering Sea to Panamd; north Japan 
Sea to Japan; west coast South America (ChUe), Straits of Magellan, 
Falkland Islands. In low water to 110 fathoms. 

Family Sternaspidae 

Body very short, grublike, aberrant; segments few, uniramous or 
achaetous, without parapodial lobes (fig. 35, a). Prostomium re- 
duced to small rounded tubercle, without appendages. Mouth 
rounded, subterminal, ventral to prostomium. Body divided into 
three regions: (1) anterior, with complete rings, with lateral concentric 
rows of strong acicular yeUow setae on anterior three segments 
(anterior region may be retracted into more posterior segments); 
(2) middle, with segmental divisions absent on midventral part, 
without setae visible externally; (3) posterior, with ventral horny 
shield formed by two trapezoidal plates provided on their external 
sides with radiating bundles of stiff, barbed and smooth capillary 
setae (fig. 35, b). Numerous, long, filiform anal branchiae, inserted 
on posterior end. Anus terminal. Lives in mud. 

Represented by a single species from Point Barrow (the family is 
usually considered to have a single cosmopolitan species). 

Genus Sternaspis Otto, 1821 
Stemaspis scutata (Ranzani, 1817) 

Figure 35, a, b 

Thalassema scutata Ranzani, 1817, p. 1457, pi. 11, figs. 10-13. 

Sternaspis fossor Stimpson, 1854, p. 29, pi. 2, fig. 19. — Webster and Benedict, 
1884, p. 725.— Moore, 1909b, p. 144; 1923, p. 218.— Hartman, 1944a, pp. 336, 
342, pi. 33, fig. 15.— Berkeley and Berkeley, 1952, p. 59, fig. 123. 

Sternaspis islandica Malmgren, 1867, p. 87, pi. 14, fig. 85. 

Sternaspis scutata Moore, 1903, p. 487; 1908, p. 357; 1923, p. 218.— Fauvel, 1927, 
p. 216, fig. 76, a-g; 1932, p. 213; 1933, p. 52; 1934a, p. 60.— Monro, 1930, 
p. 178.— Annenkova, 1937, p. 185; 1938, p. 196.— Hartman, 1942c, p. 102.— 
Gorbunov, 1946, p. 39.— Wesenberg-Lund, 1949, p. 345; 1950a, p. 46; 1950b, 
p. 104; 1951, p. 98.— Hartman and Reish, 1950, p. 38. 

Sternaspis scutata var. africana Monro, 1930, p. 179. 



310 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Description. — Up to 31 mm. long, 14 mm. wide. Body variable in 
shape; may be inflated at both ends, anterior region may be retracted 
into more posterior segments. Integument densely pilose, covered 
with fine filiform papillae. Anterior region of seven segments; 
segments 1 and 5-7 achaetous; segments 2-4 each with lateral semi- 
circular row of strong, short, yellow setae which are arched, pointed 
to blunt (worn down), diminishing in size toward ventral side; pair 
of long genital papillae on segment 7. Middle region of six segments, 
weU defined except for narrow midventral strip, with bundles of capil- 
lary setae embedded in body wall. Posterior region with five or more 
segments, with ventral horny trapezoidal plates showing concentric 
and radiating striae, each plate with 16 radiating bundles of stiff 
setae, 9 on lateral and 7 on posterior borders. Anal filaments or 
branchiae very numerous, long, filiform, sometimes spiralled. Colok : 
In alcohol: Without color or brownish, with rusty-red ventral shield. 

New records. — Arctic Alaska: Off Point Barrow base, up to 5 
miles from shore, 1.7-24.7 fms., on bottoms of mud and gravel with 
mud (3 stations, 5 specimens). Bering Sea: 63°37' N., 165°19' W., 
12 fms., Stoney, 1884; off mouth Yukon Eiver, 3 fms., E. W. Nelson, 
1877. Alaska: Iluiliuk Harbor, Unalaska, Dall, 1871; Albatross 
Sta. 3340, 55°26' N., 155°26' W., 695 fms., and Sta. 3330, 54° N., 
166°53' W., 351 fms., 1890. Chiachi Islands, 20 fms., 1874; off 
Round Island, Coal Harbor, Unga Island, 6-8 fms., 1872; Eastern 
Harbor, Sitka, 15 fms., gravelly-mud, 1874; all collected by Dall. 
West Coast North America: Washington and Puget Sounds, 
10-111 fms., mud, Pettibone. East Coast North America: Nova 
Scotia, Maine, Massachusetts, 5-102 fms., U. S. Fish Commission. 

Distribution. — Scattered records in the Arctic : Siberian and Alaskan 
Arctic, Greenland. Also Iceland to English Channel, Mediterranean, 
Adriatic, Gulf of Oman, Persian Gulf; Nova Scotia to Massachusetts; 
Bering Sea to Panamd; north Japan Sea to Japan, China. Southern 
latitudes: South Shetlands, Palmer Archipelago, South Orkneys, 
South Georgia, west Africa, Indian Ocean, Australia, New Zealand. 
In 1.7-766 fathoms. 

Family Pectinariidae (Amphictenidae) 

Body short, conical, with segments few in number. Prostomium 
indistinct, fused with buccal segment forming a truncate anterior end 
provided with two bundles of paleae (}&Tge, flattened, golden setae) 
in a horizontal row directed obliquely anteriorly, forming an operculum 
for the tube or as a fork for digging in sand or mud ; ventral to paleae, 
an antennular or tentacular membrane bordered or not with short to 
long filiform papillae, surrounding numerous prehensile oral tentacles 



MARINE POLYCHAETE WORMS — PETTIBONE 311 

not retractile in mouth; dorsal and posterior to paleae a semilunar 
flat or concave cephalic plate limited posteriorly by an entire or 
crenulate rim; two pairs tentacular cirri with first pair lateral to base 
of paleae, second pair lateral to rim of cephalic plate (fig. 35, c, d). 
Body divided into three regions: (1) anterior thoracic, including two 
achaetous segments each with a pair of lateral pectinate branchiae, 
three notopodial uniramous setigers lacking neuropodia; ventral side 
anterior region with thick glandular cushions ; (2) median abdominal, 
with biramous segments (except last few may have notosetae only), 
notopodia with setae capillary, limbate, smooth or finely den tided; 
neuropodia in form of wide pinnules bearing single row of pectiniform 
uncini; (3) posterior anal plaque or scapha, small, foliaceous, concave 
dorsally, folded under abdomen, with rudimentary achaetous segments 
except for the first segment which has a series of acicular, more or 
less recurved setae (the scaphal hooks); an oval or filiform ligule 
above the anus (fig. 35, g, h). Tube free, conical, rigid, fragile, open 
at both ends, straight or arcuate, formed of a single layer of cemented 
sand grains or shells, and liued by a membrane. Live in sand or mud, 
with anterior and larger end of tube directed below. May secrete 
mucus in abundance. 

Represented by a single genus (and subgenus) and two species 
from Point Barrow. 

Genus Pectinaria Laxnark, 1818 

Subgenus Cistenides Malmgren, 1865 

Both species have the tube sHghtly arched, lined with rusty- 
colored membrane. Tentacular membrane free, the lateral portions 
not fused to paleal segment, with 30-45 filiform to clavate marginal 
papillae. Oral tentacles short, thick, grooved. Dorsal rim of flat 
paleal segment entire, not den tided (fig. 35, d). Usually 17 segments 
with capillary notosetae (15-17, the last two segments may have the 
notosetae few in number or lacking), of which 12 (beginning on setiger 
4) have elongate flattened pinnules bearing single rows of neuropodial 
uncini; pectiniform uncini thin, flat, with 3-4 major teeth and 4-5 
small ones above basal process (fig. 35, /). Anal plaque distinctly 
separated from abdomen by constriction, suboval, festooned laterally 
(rudimxcntary parapodia) ; basally with 7-1 1 pairs short, heavy scaphal 
hooks (fig. 35, g). 

Key to the species of Pectinaria (Cistenides) from Point Barrow 

1. Cephalic paleae usually 7-9 pairs (7-13), heavy, with blunt tips (or very short 

sharp tips; fig. 35, i, f) P. (C.) granulata 

Cephalic paleae usually 11-14 pairs (9-15), long, tapered to slender pointed 
tips (fig. 35, e) P. (C.) hyperborea 



312 PROCEEDINGS OF THE NATIONAL MUSEUM voL.m 

Pectinaria iCistenides) granulata (Linne, 1767) 

Figure 35, i-k 

Sabella granulata Linn6, 1767, p. 1268. 

Cistenides granulata Malmgren, 1865, p. 359. — Webster and Benedict, 1887, p. 

747.— Chamberlin, 1920, p. 25.— Berkeley and Berkeley, 1942, p. 200.— 

Hartman, 1944a, pp. 335, 342.— Wesenberg-Lund, 1950b, p. 105; 1951, p. 99. 
Pectinaria brevicoma Johnson, 1901, p. 423, pi. 15, figs. 151-156. — Not Moore, 

1923, p. 216 (= P. calif orniensis Hartman; in USNM). 
Pectinaria (Cistenides) granulata Hessle, 1917, p. 77. — Nilsson, 1928, p. 28, fig. 

8.— Annenkova, 1937, p. 186; 1938, p. 198.— Treadwell, 1937, p. 32. 
Cistenides brevicoma Hartman, 1941b, p. 331, pi. 50, figs. 13, 14, 16; pi. 52, fig. 

23; 1944b, p. 268. 
Pectinaria (Cistenides) brevicoma Berkeley and Berkeley, 1952, p. 106, figs. 218, 

219. 

Description. — Body 20-52 mm, long, 6-8 mm. wide. Tube 38-52 
mm. long, 7-8 mm. in greatest diameter, of rather coarse sand grains, 
nearly uniform in size. Paleae usually 8 or 9 pairs (7-13), short, 
heavy, with tips blunt (may have short, sharp tips). Scaphal hooks 
strongly hooked, with a distract shoulder. Color: In life: Colorless, 
transparent, with reddish-orange internal organs, red branchiae, 
golden paleae. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, stony; Point Barrow base, washed ashore; off Point 
Barrow base, up to 12.1 miles from shore, 3.7-123.5 fms., on bottoms 
of mud, stones, mass of worm tubes, and various combinations of 
mud, sand, gravel, rocks, stones, large perforated rocks, shells (17 
stations, 53 specimens); 10 miles west Point FrankHn, 13.5 fms., 
sand, Point Barrow Expedition, 1883. Kamchatka: Petropavlovsk, 
Grebintzky, 1883. Bering Sea: Atka Island, Aleutians, Turner, 1879; 
Bering Straits, 13 fms., DaU, 1880. Alaska: Chichagof Harbor, Attn 
Island, 5-7 fms., 1873; Coal Harbor, Unga Island, 1872; and Port 
Etches, 12-18 fms., 1874; all collected by Dall. Kodiak, Fisher; 
Wrangel, Jones. Washington: Puget Sound, 39-83 fms., mud, rocks, 
shells, Pettibone. Canadian Arctic: Kneeland Bay, Frobisher 
Bay, Baffin Island, and off Daniels Island, Newell Sound, Frobisher 
Bay, 10-30 fms., Bartlett, 1942. Northwest Greenland: Off 
Conical Eock, 76° N., 67°30' W., 20-40 fms., 1938; between Parker 
Snow Bay and Conical Rock, 25-45 fms., 1940; west side Wolstemholm 
Island, 12 fms., 1940; all collected by Bartlett. East Coast North 
America: Off Labrador, 6-12 fms., Bliie Dolphin Expeditions, 1949, 
1950, 1951; off New Brunswick, Nova Scotia, Maine, New Hampshire, 
Massachusetts, Long Island Sound, 5-190 fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Novaya Zemlya. Also 



MARINE POLYCHAETE WORMS — PETTIBONE 



313 







J) 



K 



y 



J 



Figure 35. — Sternaspidae: a, Sternaspis scutata, ventral yiew anterior end; h, same, ventral 
view posterior end. Pectinariidae: c, Pectinaria hyperhorea, ventral view anterior end; 
d, same, dorsal view anterior end; e, same, palea;/, same, pectiniform uncinus; g, same, 
dorsal view posterior anal plaque or scapha; h, same, scaphal seta; i, Pectinaria granulata, 
cephalic paleae; /, same, cephalic paleae, with short mucronate tips; ^, same, scaphal 
hook. (For explanation of symbols, see p. 210.) 



314 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Iceland, Faroes; Labrador to Long Island Sound; Bering Sea to western 
Mexico; north Japan Sea. In low water to 190 fathoms. 

Pectinaria (Cistenides) hyperborea (Malmgren, 1865) 

FiGUKE 35, c-h 

Cistenides hyperborea Malmgren, 1865, p. 360, pi. 18, fig. 40. — Moore, 1903, p. 479. — 
Fauvel, 1914, p. 277, pi. 26, figs. 27, 28.— Augener, 1928, p. 775.— Berkeley 
and Berkeley, 1942, p. 201. — Wesenberg-Lund, 1950a, p. 46; 1950b, p. 106; 
1951, p. 100. 

Pectinaria (Cistenides) hyperborea Hessle, 1917, p. 76. — Nilsson, 1928, p. 31, fig. 
9.— Gustafson, 1936, p. 8.— Okuda, 1937b, p. 56, fig. 5; pi. 2, fig. F.— Annen- 
kova, 1937, p. 186; 1938, p. 197.— Zatsepin, 1948, p. 147, pi. 37, fig. 4. 

Description. — Body 28-55 nun. long, 6-12 mm. wide. Tube 45-72 
mm. long, 7-10 mm. in greatest diameter, formed of fine sand grains 
plus some coarser ones. Paleae usually 11-14 pairs (9-15), long, 
tapered to slender tips. Scaphal hooks taper gradually, without 
distinct shoulder. 

New records. — Arctic Alaska: Off Point Barrow base, up to 15 
miles from shore, 6-123.5 fms., on bottoms of mud, mass of worm 
tubes, and various combinations of mud, gravel, stones, rocks, with 
shells, worm tubes; from fish trap (8 stations, 19 specimens). Bering 
Sea: Albatross Sta. 3610, 55°58' N., 167°16' W., 75 fms., mud, 1895. 
Alaska: Albatross Sta. 4244, Kasaan Bay, Prince of Wales Island, 
50-54 fms. West Greenland: Godhavn, U.S.S. Alert, 1884. East 
Coast North America: Off Labrador, 8-125 fms.. Blue Dolphin 
Expeditions, 1949, 1950, 1951; off Nova Scotia, Maine, Massachusetts, 
16-65 fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Spitsbergen, Novaya Zemlya, Kara Sea. 
Also Iceland, Norway to Danish waters. North Sea; Labrador to 
Massachusetts; Bering Sea to Alaska; north Japan Sea to Japan. 
In 1.5-379 fathoms. 

Family Ampharetidae 

Prostomium (or tentacular membrane) more or less distinctly 
trilobed, with numerous filiform oral tentacles, smooth or pinnate, 
retractile in mouth (fig. 36, a, b, e). Branchiae filiform or subulate 
(rarely pinnate), two to four pairs, inserted on dorsal part of first 
setigerous segments. First few segments achaetous, with or without 
special group of setae or paleae anterior to branchiae. Body divided 
into two distinct regions: (1) thoracic; conical notopodia with smooth, 
limbate capillary setae and, beginning on setigers 3 or 4, with neuro- 
podial flattened pinnules bearing a row of pectiniform uncini; (2) 
abdominal; uncinigerous pinnules only. With or without dorsal. 



MARINE POLYCHAETE WORMS — PETTIBONE 315 

ventral or anal cirri. Tube membranous, covered with mud or ag- 
glutinated foreign bodies. 

Eepresented by two genera and four species at Point Barrow. 
Both genera have prostomia trilobed, without projecting glandular 
crests; oral tentacles numerous, pinnate; without large dorsal hooks 
posterior to branchiae; branchiae four pairs, filiform or subulate, 
arising on or near a distinct transverse dorsal ridge; thoracic setigers 
14; uncinigerous pinnules begin on setiger 3; notopodia without 
claviform cirri. 

Key to the genera of Ampharetidae from Point Barrow 

1. With paleae, lateral group of golden setae anterior to branchiae (fig. 36, 

o, b) Ampharete (p. 315) 

Without paleae Asabellides (p. 318) 

Genus Ampharete Malmgren, 1865 

All three species have prostomium distinctly trilobed; median lobe 
with pair of basal eyes, lateral lobes encircling median lobe posteriorly 
(fig. 36, a). First two segments achaetous; first or buccal segment 
extended anteroventrally forming large lower lip. Segment 3 or 
paleal segment with group of paleae an*anged in semicircle just ante- 
rior and lateral to branchiae. Abdominal segments with neuropodial 
cirri dorsally on uncinigerous pinnules poorly developed (short, 
rounded to conical) to well developed (cirrif orm, longer than pinnule) . 

Key to the species of Ampharete from Point Barrow 

1. First 2 abdominal segments with enlarged, padlike, achaetous notopodial 

lobes A. vega 

Without enlarged notopodial lobes on first few abdominal segments 2 

2. Paleae slender, taper gradually. Abdominal setigers 12, usually with promi- 

nent neuropodial cirri. Anal cirri numerous (fig. 36, c) A. acutifrons 

Paleae taper rather abruptly, with short mucronate tips. Abdominal setigers 
16-17, with neuropodial cirri inconspicuous. Anal cirri 2 A. goesi 

Ampharete vega (Wiren, 1883) 

Figure 36, a 

Amphideis vega Wir6n, 1883, p. 415, pi. 32, figs. 3, 4. 

Ampharete vega Hessle, 1917, p. 99. — Augener, 1928, p. 778. — Annenkova, 1929, 
p. 493.— Zatsepin, 1948, p. 150. 

Description. — Length 16 mm., width 1.5 mm. (up to 50 mm. long, 
4 mm. wide — Wiren, 1883). Paleae slender, tapering gradually to 
fine tips, 10-13 in each group. Eight branchiae form continuous 
transverse group, arising from low fold. Abdominal segments up to 
27-28 (16 in Point Barrow specimen; complete?). First two abdom- 

261112—54 8 



316 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

inal segments with enlarged, rounded, padlike, achaetous notopodial 
lobes. Neuropodial cirri poorly developed or lacking. Pygidium with 
several small papillae? Colorless in alcohol. Tube thin, membra- 
nous, with sand and debris, rather straggly. 

New records. — Arctic Alaska: West side Elson Lagoon near en- 
trance to small lagoon to west, near Point Barrow (rather brackish 
water), 1.2 fms. (1 station, 1 specimen). 

Distribution. — Scattered records in the Arctic : Siberian and Alaskan 
Arctic, Spitsbergen, Barents Sea, Kara Sea, Laptev Sea. In L2-11 
fathoms. 

Ampharete acutifrons (Grube, 1860) 

Figure 36, b-d 

Amphideis acutifrons Grube, 1860, p. 109, pi. 5, fig. 6. 

Ampharete grubei Malmgren, 1865, p. 363, pi. 19, fig. 44. — Eliason, 1920, p. 70. — 

Fauvel, 1927, p. 227, fig. 79, a-p.— Augener, 1928, p. 776.— Thorson, 1946, 

p. 121, fig. 66.— Hartman, 1948, p. 43. 
Ampharete cirrata Webster and Benedict, 1887, p. 747, pi. 8, figs. 110-112. 
Ampharete trilobata Webster and Benedict, 1887, p. 747. 
Ampharete acutifrons Hessle, 1917, p. 96. — Annenkova, 1937, p. 188; 1938, p. 

200.— Zatsepin, 1948, p. 150, pi. 37, fig. 8.— Wesenberg-Lund, 1950a, p. 47; 

1950b, p. 109; 1951, p. 102. 

Description. — Length 45 mm., width 6 mm. (up to 56 mm. long, 
8 mm. wide — Wiren, 1883). Paleae slender, taper gradually to long, 
delicate tips, may be somewhat curved, 10-30 in each group. Bran- 
chiae in two groups separated middorsally, each group with three 
branchiae in transverse line and one slightly more posterior. Abdom- 
inal segments 12. With neuropodial cirri small, conical to quite 
long, longer than pinnule (weU developed on Point Barrow specimens). 
Pygidium a short ring bearing a circle of numerous subequal anal 
cirri (2 lateral ones may be longer). Colorless in alcohol. Tube 
membranous, covered with soft, gray debris (or agglutinated mud). 

Remarks. — The types of Ampharete cirrata Webster and Benedict 
from Eastport, Maine, were examined. This species was separated 
on the basis of the relative development of the neuropodial cirri, 
which appears to be somewhat variable. The types of A. trilobata 
Webster and Benedict, also from Eastport, were not available, but 
this species also, according to the original description, was differen- 
tiated by the relative development of the cirri. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow; Point Barrow base, washed ashore; off Point Barrow 
base, up to 6 miles from shore, 3.7-49 fms., on bottoms of mud, and 
various combinations of mud, pebbles, gravel, stones, rocks, and 



MARINE POLYCHAETE WORMS — PETTIBONE 317 

shells (10 stations, 17 specimens). West Coast North America: 
Washington and Puget Sounds, 80 fms., Pettibone. East Coast 
North America: Off Labrador, 8 fms,, Blue Dolphin Expedition, 
1949. 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Jan Mayen, Spitsbergen, Novaya Zem- 
lya. Also Iceland, Swedish west coast to France, Mediterranean; 
Labrador to Maine; Bering Sea to southern California; north Japan 
Sea. In 1-1,333 fathoms. 

Ampharete goesi Malmgren, 1865 

Ampharete goesi Malmgren, 1865, p. 364, pi. 19, fig. 45. — Hessle, 1917, p. 91. — 
Augener, 1928, p. 778.— Annenkova, 1929, p. 492, pi, 38, fig, 37; 1937, p. 188; 
1938, p. 200.— Zatsepin, 1948, p. 150, pi. 38, fig. 10.— Wesenberg-Lund, 
1950a, p. 47; 1950b, p. 110; 1951, p. 102.— Berkeley and Berkeley, 1952, p. 
66, fig. 136. 

Ampharete goesi subsp. braznikovi Annenkova, 1929, p. 492, pi. 38, fig. 44. 

Description. — Length 35 mm,, width 6 mm. (up to 50 mm. long, 7 
mm. wide — Malmgren, 1865). Paleae rather stout, taper abruptly, 
ending in short acuminate tips, 14-21 in each group. Branchiae in 
two groups separated middorsally, each group with three branchiae 
in transverse line. Abdominal segments 17 (stem form and Point 
Barrow specimens) or 16 (subsp. braznikovi and Washington Sound 
specimens). Neuropodial cirri inconspicuous, low, rounded. Pygid- 
ium with two long anal cirri. Color: In life: Body reddish orange, 
branchiae green. In alcohol: Colorless. Tube: Membranous, rather 
thick, with debris including bits of shells, sea urchins tests, and 
foraminiferans, very straggly. 

New records. — Arctic Alaska: Off Point Barrow base, up to 15 
miles from shore, 20.3-123.5 fms., on bottoms of mass of worm tubes 
and various combinations of mud, pebbles, gravel, stones, rocks, shells, 
and worm tubes (11 stations, 20 specimens). West Coast North 
America: Washington Sound, Pettibone. 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Spitsbergen, Barents Sea, White Sea, 
Novaya Zemlya. Also Iceland; British Columbia to Washington; 
Okhotsk Sea to north Japan Sea. In 10.5-353 fathoms. 

Ampharete sp. (Young) 

Two very small specimens, 12 mm. long, were found on bottom of 
rocks with bryozoans and hydroids in 17 fathoms September 9, 1948. 
Prostomium 3-lobed, with four eye spots, bases of four pairs branchiae, 
14 thoracic segments, 10 abdominal segments plus a growing zone. 



318 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

GerkusAsabellides Annenkova, 1929 

Includes Pseudosahellides Berkeley and Berkeley, 1943. 

Asabellides sibirica (Wiren, 1883) 

Figure 36, e 

Sabellides sibirica Wir6n, 1883, p. 418. 

Asabellides orientalis Annenkova, 1929, p. 494, pi. 38, figs. 50, 51, pi. 39, figs. 

60-65; 1937, p. 188. 
Asabellides sibirica Annenkova, 1938, p. 201. 
Pseudosabellides lineata Berkeley and Berkeley, 1943, p. 131; 1944, p. 3; 1952, 

p. 71, fig. 147. 

Description. — Body long, slender, 38 mm. long, 1.5 mm. wide. 
Prostomium trilobed with median lobe wider and flared anteriorly, 
lateral lobes diagonal. Segments 1 and 2 fused laterally and ventrally, 
forming large lower lip; dorsally, appear as two short segments more 
or less hidden by dorsal branchial fold. Segments 3 and 4 more or less 
fused ventrally and laterally; dorsally, segment 3 without paleae but 
with branchiae; segment 4 with first notopodial lobes and, middorsally, 
a fused nephridial area wath pair of papillae posterior to branchial 
bases. Branchiae slender, fihform, emerge from prominent ridge, six 
in transverse row, two slightly posterior. Abdominal segments 23 
(14-23). Neuropodial cirri smaU, short, rounded (Point Barrow 
specimens) to long, slender, cirriform. Pygidium with few small cirri 
and pair of long lateral ones. Color: In life: Salmon below, darker 
above, branchiae green, oral tentacles pink. In alcohol: Colorless. 

New records. — Arctic Alaska: Off Point Barrow base, 5 mUes from 
shore, 11.7 fms., on bottom of mud (1 station, 3 specimens). 

Distribution. — Scattered records in the Arctic: Siberian, Alaskan, 
and Canadian Arctic. Also Hudson Bay; Bering Sea to British 
Columbia; Okhotsk Sea to north Japan Sea. In 4-106 fathoms. 

Family Terebellidae 

Body divided into two regions: (1) thoracic; more or less inflated, 
with dorsal bimdles of capillary setae and ventral uncinigerous tori 
(sometimes acicular crotchets, sometimes absent), often with ventral 
glandular shields; (2) abdominal; elongated, more or less tapering, 
most often lacking dorsal setae and bearing only uncinigerous pinnules. 
Prostomium of variable form and dimensions, fused with buccal seg- 
ment, forming a lower lip, a transverse dorsal cephalic ridge with or 
without numerous eye-spots, a tentacular membrane with numerous 
filiform grooved tentacles not retractile in mouth, a semicircular or 
folded upper lip (fig. 36,/). Branchiae 1-3 pairs, filiform or arbores- 
cent, or single, quadrilobed, pectinate, on anterior segments, or 



MARINE POLYCHAETE WORMS — PETTIBONE 



319 




Figure 36. — A.mph.arttid&e: a, Ampharetevega, dorsal view anterior end; b, Ampharete acuti- 
frons, dorsal view anterior end; c, same, lateral view posterior end; d, same, palea; i, 
Asabellides sibirica, dorsal view anterior end. Terebellidae: /, lateral view anterior end 
of terebellid; g, Amphitriu cirraia, lateral view anterior end; h, same, notoseta; i, Nicolea 
venustula, lateral view anterior end of male; ;, same, thoracic uncini; k, Pista maculata, 
lateral view anterior end, tentacles missing; /, same, uncinus with long manubrium. (For 
explanation of symbols, see p. 210.) 



320 PROCEEDINGS OF THE NATIONAL MUSEUM tol. 103 

branchiae lacking. Without paleae. Tube usually membranous, 
encrusted with sand, debris of shells, algae, etc. 

Represented by 10 genera and 11 species at Point Barrow. 

Key to the genera of TerebeUidae from Point Barrow 

1. Uncini in single rows on first 6 uncinigerous segments, then in 2 rows (alter- 

nating, intercogging or opposite) on certain number of following segments. 
With distinct thoracic ventral shields (single, not paired) . Notosetae begin 
on segment 4; uncini begin on segments 5 or 6 (setigers 2 or 3), Fused 
prostomium and buccal segment forming a prominent rectangular lower lip, 
a transverse semicircular cephalic ridge (with or without numerous eye-spots 
along basal groove), flared semicircular upper lip, semicircular depressed 
tentacular region between upper lip and cephalic ridge with numerous, 

filiform, grooved tentacles (fig. 36,/) 2 

Uncini in single rows only (or with crotchets on thoracic segments) 7 

2. With 1-3 pairs branchiae dorsally on segments 2-4 (fig. 36, g, i, k). Uncini 

begin on segment 5 (setiger 2) 3 

Without branchiae. With lateral lobes on anterior segments (fig. 37, o) 5 

3. Notosetae with tips finely denticled (fig. 36, h). Cephalic ridge usually 

without eye-spots. With lateral lobes on anterior segments (fig. 36, g). 

Amphitrite (p. 321) 
Notosetae with tips smooth. Cephalic ridge usually with eye-spots 4 

4. Without lateral lobes on anterior segments. Uncini of anterior segments 

without long manubrium (fig. 36, j). Branchiae 2 pairs, branched dichoto- 
mously (fig. 36, t) Nicolea (p. 322) 

With large lateral lobes on buccal segment, enclosing prostomium and con- 
nected ventrally (fig. 36, k). Uncini of first few segments with long manu- 
brium (fig. 36, I). Branchiae 1-3 pairs, arborescent, with main trunk well 

marked Pista (p. 323) 

6. Uncini begin on segment 6 (setiger 3). Notosetae of 2 kinds, smooth and 
denticled Proclea (p. 325) 

Uncini begin on segment 5 (setiger 2). Notosetae with tips smooth 6 

6. Uncini in single rows on first 6 uncinigerous segments, in double rows on next 

10. Nephridial papillae of segment 3 not elongated. Notosetae broadly 

bilimbate below whipUke tip (fig. 37, b) Leaena (p. 325) 

Uncini in single rows on first 6 uncinigerous segments, in double rows on next 
8. Nephridial papillae of segment 3 (anterior to first setiger) elongated. 
Notosetae weakly limbate on one side along most of its length (fig. 37, c). 

Lanassa (p. 326) 

7. Uncini all short, avicular (fig. 37, d) 8 

Uncini of 2 types; thoracic aciculiform, with long manubrium (fig. 37, h); 

abdominal avicular (fig. 37, i). Without ventral shields 9 

8. Branchiae 2-3 pairs, cirriform, in transverse rows. Notosetae begin on 

segment 3, continuing on large number of segments; uncini begin on segment 
5 (setiger 3). Ventral shields usually slightly distinct. Tentacular lobe 

not especially enlarged Thelepus (p. 327) 

Without branchiae. Notosetae begin on segments 2 or 3; uncini begin on 
variable number of segments, may be absent from thoracic region. With 
ventral shields, first unpaired, rest paired, more or less separated in median 
line (fig. 37, /). Tentacular membrane large, trilobed or undulating, 
bearing very numerous tentacles (fig. 37, e-/), Polycirrus (p. 328) 



MARINE POLYCHAETE WORMS — PETTIBONE 321 

Branchiae 2-3 pairs, single, filiform (fig. 37, g). Notosetae and uncini 
(crotchets) both begin on segment 6. Tentacular lobe folded, with very- 
numerous thin, cylindrical, and short, grooved tentacles, 

Trichobranchus (p. 329) 

Branchiae single, consisting of large cylindrical trunk and 4 pectinate lobes 
(fig. 37, j). Notosetae begin on segment 3, uncini begin on segment 8 
(setiger 6). Tentacular lobe large, folded, with numerous tentacles of a 
single kind Terebellides (p. 330) 



Genus Amphitrite O. F. Muller, 1771 

Both species have the body inflated anteriorly, decidedly atten- 
uated posteriorly. Cephalic ridge without eye-spots. Branchiae 
three pairs, on segments 2-4. Notosetae begin on segment 4; uncini 
begin on segment 5, in single rows on first six, in double rows on rest 
of thoracic setigers, in single rows on abdominal segments on project- 
ing pinnules. Pygidium a crenulate ring. 

Key to the species of Amphitrite from Point Barrow 

1. Branchiae cirriform, a tuft of simple filaments from short, sUghtly swollen 

common base (fig. 36, g). Thoracic setigers 17 A. cirrata 

Branchiae branched. Thoracic setigers 19 A. groenlandica 

Amphitrite cirrata Muller, 1776 

Figure 36, g, h 

Amphitrite cirrata O. F. Mtiller, 1776, p. 216.— Malmgren, 1865, p. 375, pi. 21, 
fig. 53.— Webster and Benedict, 1887, p. 748.— Hessle, 1917, p. 185.— 
Chamberhn, 1920, p. 22.— Eliason, 1920, p. 74.— Fauvel, 1927, p. 251, 
fig. 86, i-o.— Augener, 1928, p. 785.— Okuda, 1938b, p. 102.— Hartman, 
1944a, pp. 334, 342; 1948, p. 43.— Thorson, 1946, p. 127, figs. 71, 72.— 
Gorbunov, 1946, p. 39.— Zatsepin, 1948, p. 156, pi. 38, fig. 12.— Wesenberg- 
Lund, 1950a, p. 50; 1950b, p. 116; 1951, p. 106.— Berkeley and Berkeley, 
1952, p. 86, fig. 175. 

Amphitrite radiata Moore, 1908, p. 350; 1923, p. 193. 

Description. — Length up to 200 mm., width 10 mm. About 12 
ventral shields. Branchiae 3 pairs, cirriform, a tuft of simple fila- 
ments from a short common base. Each of thi-ee branchial segments 
(segments 2-4) with pair of lateral lobes. Nephridial papillae 7 
pairs; on segment 3, prominent papillae ventral to second branchial 
pair; on segments 6-11, rather inconspicuous papillae between noto- 
podia and neuropodia. Thoracic setigers 17. Color: In alcohol: 
Keddish brown. Tube: Mostly of mud, with scattered small pebbles 
and debris, soft, breaking easily. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon; off 
Point Barrow base, up to 7 miles from shore, 18.3-70 fms., on various 
combinations of mud, sand, gravel, stones, rocks, large perforated 
rocks, shells (8 stations, 14 specimens). West Coast North 



322 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

America: Wrangel, Alaska, Jones; Puget Sound, low tide, Pettibone. 
West Greenland: Vaigat, Disko Island, Bartlett, 1937. East 
Coast North America:] Bay of Fundy, Maine, Massachusetts, 
14-llO^fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic : Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitzbergen, Franz Josef Land, 
White Sea, Novaya Zemlya, Kara Sea. Also Iceland, Faroes, Nor- 
way to British Isles, Azores, Mediterranean, Adriatic; Bay of Fundy 
to Massachusetts; Bering Sea to southern California; Japan. In low 
water to 1,528 fathoms. 

Amphitrite groenlandica Malmgren, 1865 

Amphitrite groenlandica Malmgren, 1865, p. 376, pi. 21, fig. 52. — Verrill, 1881, 
pp. 305, 310.— Hessle, 1917, p. 181.— Fauvel, 1927, p. 250, fig. 86, a-c— 
Augener, 1928, p. 787.— Amaenkova, 1937, p. 192; 1938, p. 206.— Gorbunov, 
1946, p. 39.— Zatsepin, 1948, p. 157, pi. 38, fig. 14.— Wesenberg-Lund, 
1950a, p. 51; 1950b, p. 117; 1951, p. 107. 

Neoamphitrite robusta Hartman, 1948, p. 44 (not Amphitrite robusta J ohnaon, 
1901). 

Description. — ^Length up to 190 mm., width 9 mm. (a single, much 
smaller specimen from Point Barrow). About 14 ventral shields. 
Branchiae 3 pairs, branched dichotomously 3-4 times, with main 
trunk short, thick. Lateral lobes of first two branchial segments well 
developed, continuing ventrally; small lateral lobes on third branchial 
segment. Nephridial papillae 12 pairs on segments 3-14, the first 
four with more prominent papUlae. Thoracic setigers 19. Colorless 
in alcohol. Tube of mud. 

Remarks. — A. robusta Johnson is very close to A. groenlandica; 
it differs in having 17 thoracic setigers instead of 19. 

New records. — Arctic Alaska: Off Point Barrow base, 5 miles from 
shore, 49 fms., on bottom of rocks, stones, gravel (1 station, 1 speci- 
men). Southwestern Alaska: Round Island, Coal Harbor, Unga 
Island, 8-9 fms., Dall, 1872. 

Distribution. — Scattered records in the Arctic: Siberian and Alaskan 
Arctic, West Greenland, Spitsbergen, Novaya Zemlya. Also Iceland, 
Scandinavian coast to North Sea, Ireland; Maine; southwestern 
Alaska; Okhotsk Sea to north Japan Sea. In 7 to 440 fathoms. 

Genus Nicolea Malmgren, 1865 
Nicolea venustula (Montagu, 1818) 

Figure 36, i, j 

Terebella venustula Montagu, 1818, p. 344, pi. 13, fig. 2. 

Nicolea arctica Malmgren, 1865, p. 381, pi. 24, figs. 66, 67. 

Nicolea zostericola Malmgren, 1865, p. 381, pi. 26, fig. 76. — Webster and Benedict, 
1887, p. 749.— Moore, 1909b, p. 141.— Fauvel, 1927, p. 261, fig. 90, g-n.— 
Annenkova, 1934, p. 322; 1937, p. 191; 1938, p. 205.— Gustafson, 1936, p. 



MARINE POLYCHAETE WORMS — PETTIBONE 323 

9.— Berkeley and Berkeley, 1943, p. 130; 1952, p. 87, figs. 177, 178.— Gor- 

bunov, 1946, p. 39.— Hartman, 1948, p. 44. 
Nicolea simplex Verrill, 1873, p. 613; 1881, p. 302, pi. 10, fig. 1.— Sumner, Osburn, 

and Cole, 1913, p. 627. 
Nicolea venustula Ehlers, 1913, p. 559. — Hessle, 1917, p. 171. — Chamberlin, 1920, 

p. 22.— Eliason, 1920, p. 73.— Fauvel, 1927, p. 260, fig. 90, a-f ; 1934a, p. 68.— 

Augener, 1928, p. 788.— Thorson, 1946, p. 126.— Zatsepin, 1948, p. 156, pi. 

38, fig. 11.— Wesenberg-Lund, 1950a, p. 51; 1950b, p. 118; 1951, p. 109. 

Description. — Length 15-48 mm., width 2-5 mm., segments 30-46 
(up to 70 mm. long — Wiren, 1883). Ventral shields about 14. 
Cephalic ridge with numerous dark eye-spots. Branchiae two pairs, 
on segments 2-3, branched dichotomously 3-6 times, with very short 
main stem, first pair much larger than second. Without lateral lobes 
on branchial segments. Nephridial papillae on segments 3 (slightly 
posterior to pair of second branchiae), 6, and 7; small in female; in 
male, papillae on segments 6 and 7 long, cylindrical. Thoracic seti- 
gers 15 (14-18, according to Hessle). Uncini begin on segment 5 
(setigerous segment 2), in single rows on first six, in two alternating 
rows on rest of thoracic segments, in single rows on prominent pro- 
jecting abdominal pinnules. Pygidium crenulate. Colorless or 
slightly brownish in alcohol. Females with large yolky eggs inside 
body (Point Barrow, August 21, and September 15, 1948; August 8, 
1949). Tube with tough, translucent membranous lining, with small 
pebbles of various sizes, shell debris, foraminiferans, etc. 

New records. — Arctic Alaska: Off Point Barrow base, up to 12.1 
miles from shore, 13.3-123.5 fms., on bottoms of mud, stones, mass of 
worm tubes, and various combinations of mud, sand, gravel, pebbles, 
stones, rocks, large perforated rocks, with bryozoans, hydroids, shells 
(24 stations, 79 specimens). Alaska: Security Bay, Jones. Cana- 
dian Arctic: Dobbin Bay, E. Ellesmere Island, 79°36' N., 73°35' 
W., 17 fms., Littlewood, 1950. East Coast North America: Off 
Maine, Massachusetts, Rhode Island, Connecticut, shore to 63 
fathoms and surface (young), U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Jan Mayen, Spitsbergen, Franz 
Josef Land, Novaya Zemlya, Kara Sea. Also Iceland, Faroes, Nor- 
way to France, Mediterranean, Adriatic, Red Sea; Hudson Bay to 
Connecticut; Bering Sea to western Canada; north Japan Sea; South 
Africa. In low water to 472 fathoms. 

Grenus Pista Malmgren, 1865 
Pista maculata (Dalyell, 1853) 

Figure 36, k, I 
Terebella maculata Dalyell, 1853, p. 203, pi. 28, figs. 10-14, 19. 
Scione lobata Malmgren, 1865, p. 383, pi. 23, fig. 62.— Verrill, 1881, pp. 305, 310.— 

Webster and Benedict, 1887, p. 749. — Augener, 1928, p. 789. — Friedrich, 

1939, p. 127. 



324 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Pista maculata Hessle, 1917, p. 161, pi. 3, fig. 4. — Fauvel, 1927, p. 263, fig. 91, 
a-h. — Berkeley and Berkeley, 1943, p. 130, — Gorbunov, 1946, p. 39. — 
Zatsepin, 1948, p. 155, pi. 38, fig. 9.— Wesenberg-Lund, 1950a, p. 52; 1950b, 
p. 119; 1951, p. 110. 

Pista groenlandica Treadwell, 1937, p. 33, figs. 14-16. 

Description. — ^Length up to 150 mm., width 6 mm. Ventral shields 
about 14. Cephalic ridge with numerous eye-spots. Single pair 
large branchiae on segment 2, with large, cylindrical main trunk, 
branched. Buccal segment with two large, rounded lateral lobes 
embracing prostomium, connected ventrally by a crest; segment 2 
short, without lateral lobes; segment 3 with large, rounded, flattened 
lateral lobes. Nephridial papillae on segments 6-7. Thoracic 
setigers 16 (one had small lobe on one side only on 17). Uncini 
begin on segment 5 (setiger 2), in single rows on first 6, in two alternat- 
ing intercogging rows on rest of thoracic segments, in single rows on 
rectangular pinnules in abdominal region. Pygidium with six or 
seven (6-12) conical papillae, arranged starlike. Color: In alcohol: 
Branchiae and tentacles dark brown or colorless. Tubes: Irregularly 
twisted, with thin transparent membranous lining covered mostly 
with fine sand grains plus few large pebbles, bits of shell, bryozoans, 
foraminiferans, algae, parts of other worm tubes. 

Remarks. — The type of P. groenlandica Treadwell from Baflin Bay 
was examined and is herein referred to P. maculata. Contrary to the 
original description, eye-spots are present on the cephalic ridge (small, 
in transverse groove, easily overlooked) . 

New records. — Arctic Alaska: Off Point Barrow base, up to 15 
miles from shore, 21-123.5 fms., on bottoms of mass of worm tubes 
and various combinations of mud, gravel, stones, rocks, large per- 
forated rocks; from crab, Hyas coarctatus (13 stations, 168 plus speci- 
mens). North Greenland: 5 miles south Cape Chalon; north 
Omenolu near North Star Bay, 17 fms., Bartlett, 1932. West 
Greenland: Oelrichs Bay, 1937; off Conical Kock, 76°3' N., 67°30' 
W., 20-40 fms., 1938; 1 mile northwest Conical Rock, 25-60 fms., 
1940; between north shores Parkers Snow Bay and Conical Rock, 
25-45 fms., 1940; aU collected by Bartlett. East Greenland: Off 
Cape Hold with Hope, 23^0 fms., Bartlett, 1939. East Coast 
North America : Off Labrador, 8-125 fms.. Blue Dolphin Expeditions, 
1949, 1950, 1951. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Baffin Bay, Greenland, Spitsbergen, Barent 
Sea, Novaya Zemlya. Also Iceland, Norway to English Channel: 
Hudson Bay to Maine; Bering Sea. In 3-1,528 fathoms. 



AIARINE POLYCHAETE WORMS — PETTIBONB 325 

Genus Proclea Saint- Joseph, 1894 
Proclea graffii (Langerhans, 1884) 

Leaena graffii Langerhans, 1884, p. 262, pi. 15, fig. 21. 

Proclea graffi Southern, 1914, p. 120.— Hessle, 1917, p. 199, fig. 53.— Fauvel, 

1927, p. 268, fig. 94, a-g.— Gorbunov, 1946, p. 39.— Zatsepin, 1948, p. 157, pi. 

38, fig. 19.— Wesenberg-Lund, 1951, p. 111. 

Description. — Length up to 42 mm., width 3 mm., segments 49. 
Ventral shields about 10. Cephalic ridge without eye-spots. With- 
out branchiae. With lateral lobes on segments 2-4, those on segments 
2 and 3 elongate, extending to ventral shields. Nephridial papillae 
on segments 3, 6-8. Thoracic setigers 16. Uncini begin on seg- 
ment 6 (setiger 3), in single rows on first six, in double rows on rest 
of thoracic segments, in single rows on projecting abdominal pinnules. 
Pygidium crenulate. Colorless in alcohol. Tube ? 

New records. — Arctic Alaska: Off Point Barrow base, up to 7.5 
miles from shore, 21-36 fms., on bottoms of rocks, stones, large per- 
forated rocks, from breaking rocks apart (3 stations, 6 specimens) . 

Distribution. — Scattered records in the Arctic : Siberian and Alaskan 
Arctic, Franz Josef Land, White Sea. Also Bering Sea to Okhotsk 
Sea; Iceland, Swedish west coast, Finland, Ireland, Madeira. In 
1-36 fathoms. 

Genus Leaena Malmgren, 1865 

Leaena abranchiata Malmgren, 1865 

Figure 37, a, b 

Leaena abranchiata Malmgren, 1865, p. 385, pi. 24, fig. 64. — not Moore, 1909b, 
p. 141 (^Lanassa venusta). — Ehlers, 1913, p. 563. — Hessle, 1917, p. 197, — 
Augener, 1928, p. 793.— Annenkova, 1937, p. 192; 1938, p. 208.— Berkeley 
and Berkeley, 1943, p. 130.— Gorbunov, 1946, p. 39.— Zatsepin, 1948, p. 157, 
pi. 38, fig. 17.— Hartman, 1948, p. 45. —Wesenberg-Lund, 1950a, p. 53; 
1950b, p. 121; 1951, p. 112. 

Leaena antarctica Mcintosh, 1885, p. 462, pi. 48, figs. 9, 10; pi. 28A, figs. 10, 11. 

Leaena abranchiata var. antarctica Hessle, 1917, p. 197. — Monro, 1930, p. 188; 
1936, p. 178. 

Description. — Length 43 mm., width 3 mm. (up to 75 mm. long, 6 
mm. wide — Malmgren, 1865). About 10 ventral shields. Cephalic 
ridge without eye-spots. Without branchiae. Lateral lobes on seg- 
ment 2 extend ventrally, those on segment 3 extend dorsally, forming 
a prominent ridge (segment preceding first setiger; ridge not so prom- 
inent in var. antarctica). Thoracic setigers 10. Uncini begin on seg- 
ment 5 (setiger 2), in single rows on first 6, in double rows on next 10, 
then in single rows on prominent abdominal pinnules. Notosetae 
widely bilimbate below whiplike tips. Colorless in alcohol. Tube : 
Of soft mud with a few bits of scattered rock (some small ones on 
tubes of Pista maculata) . 



326 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

New records. — Arctic Alaska: Off Point Barrow base, 25 fms., on 
bottoms of gravel, stones, shells (1 station, 1 specimen). East Coast 
North America: Off Labrador, 15-55 fms., mud and mud with rock, 
Blue Dolphin Expeditions, 1950, 1951. 

Distribution. — Widely distributed in the Arctic: Siberian and Alas- 
kan Arctic, Baffin Bay, Greenland, Spitsbergen, Novaya Zemlya, 
Kara Sea. Also Iceland, Faroes, Norway, Sweden, Finland; Hudson 
Bay to Labrador; southwestern Alaska; Okhotsk Sea to north Japan 
Sea. In 6-833 fathoms. Var. antarctica: Antarctic, South Georgia; 
up to 1,975 fathoms. 

Genus Lanassa M almgren, 1865 
Lanassa venustM (Malm, 1874) 

Figure 37, c 

Laphaniella venusta Malm, 1874, p. 98, pi. 1, fig. 8. 

Leaena nuda Moore, 1905b, p. 855, pi. 44, figs. 14, 15. 

Leaena abranchiata Moore, 1909b, p. 141 (not L. abranchiata Malmgren, 1865). 

Lanassa venusta Hessle, 1917, p. 205. — Eliason, 1920, p. 75. — Gustafson, 1936, 

p. 9.— Annenkova, 1937, p. 194; 1938, p. 209.— Zatsepin, 1948, p. 157. 
Lanassa venusta subsp. pacifica Annenkova, 1938, pp. 209, 230. 

Description. — Length 37-55 mm., width 3 mm., segments 54-67. 
Ventral shields about 10. Cephalic ridge without eye-spots (present in 
subsp. pacifica). Oral tentacles numerous (more than 12 as indicated 
by Malm) . Without branchiae. Lateral lobes on segments 2 and 3 
extending ventrally; small lateral lobes on segment 4. Nephridial 
papillae on segments 3, 6-8; those of segment 3 prominent (anterior 
to first setiger). Thoracic setigers 11 (12 on subsp. pacifica). Uncini 
begin on segment 5 (setiger 2), single on first six, in double rows on 
next eight, in single rows on prominent abdominal pinnules. Noto- 
setae weakly limbate (fig. 37, c). Pygidium crenulate. Females 
with large yolky eggs in body (Point Barrow, Sept. 9 and 15, 1948; 
Aug. 17, Oct. 11, 1949; Aug. 1, 1950). Color: In life: Orange, with 
white shields, tentacles tan. In alcohol: Colorless. Tube: With 
thin, transparent membranous lining, with very loose mud, bits of 
sand, rock, foraminiferans (tube of specimen from Labrador with 
great deal of loosely assembled shell debris) . 

Remarks. — The type of Leaena nuda Moore from southwestern 
Alaska was examined and found in poor condition. It has been referred 
previously to Lanassa venusta by Annenkova (1937). 

New records. — Arctic Alaska: Point Barrow base, washed ashore; 
off Point Barrow base, up to 12.1 mUes from shore, 18.3-75.5 fms., on 
bottoms of mud, stones, and various combinations of mud, sand, 
gravel, pebbles, stones, rocks, large perforated rocks, with bryozoans, 
shells (17 stations, 44 specimens). Canadian Arctic: Cove in 



MARINE POLYCHAETE WORMS — ^PETTIBONE 327 

BJQeeland Bay, Frobisher Bay, Baffin Island, 14 fms., Bartlett, 1942. 
Distribution. — Scattered records in the Arctic: Siberian, Alaskan, 
and Canadian Arctic. Also Swedish west coast, Danish waters; 
Labrador; southwestern Alaska; north Japan Sea. In 7-140.5 
fathoms. 

G^nus Thelepus Leuckart, 1849 

Thelepua cincinnatus (Fabricius, 1780) 

Figure 37, d 

Amphitrite cindnnata Fabricius, 1780, p. 286. 

Thelepus circinnaia Malmgren, 1865, p. 387, pi. 22, fig. 58. 

Thelepus cincinnatus Webster and Benedict, 1887, p. 749. — Hessle, 1917, p. 212. — 
Chamberlin, 1920, p. 23.— Eliason, 1920, p. 76.— Fauvel, 1927, p. 271, fig. 
95, i-ra; 1932, p. 233, fig. 40; 1934a, p. 69.— Augener, 1928, p. 790.— Monro, 
1930, p. 192; 1936, p. 182.— Treadwell, 1937, p. 33.— Friedrich, 1939, p. 127.— 
Hartman, 1944a, pp. 336, 343; 1952, p. 236.— Gorbunov, 1946, p. 39.— 
Zatsepin, 1948, p. 154, pi. 38, fig. 7.— Wesenberg-Lund, 1950a, p. 54; 
1950b, p. 122; 1951, p. 112. 

Thelepus hamatus Moore, 1905b, p. 856, pi. 44, figs. 16-18. — Hartman, 1948, 
p. 44. — Berkeley and Berkeley, 1952, p. 82, fig. 167. 

Description. — Up to 200 mm. long, 10 mm. wide, segments nu- 
merous. Dorsal surface rugose, with small glandular warts irregularly 
distributed and especially abundant anteriorly. Ventral shields in- 
distinct, wide, somewhat wrinkled. Buccal segment and prostomium 
forming a thick lower lip, a prominent horseshoe-shaped upper Hp, 
a semicircular cephalic ridge with numerous small eye-spots; tentacular 
area between upper lip and cephalic ridge with numerous, rather 
thick tentacles. Branchiae 2 pairs on segments 2 and 3, each formed 
of numerous, simple filaments arranged in parallel transverse rows. 
Nephridial papiUae on segments 4-7. Notosetae begin on segment 3, 
continuing on about 40 segments (or almost to posterior end). Un- 
cini begin on segment 5 (setiger 3), in single rows throughout, the 
uncinigerous tori gradually transformed into projecting rectangular 
pinnules. Pygidium with crenulate anal opening. Color: In life: 
Orange with red branchiae, tan tentacles, with whitish glandular 
areas ventrally and laterally. In alcohol: Colorless. Tube: Cylin- 
drical, twisted, with tough, transparent, membranous lining, covered 
with small pebbles of varying sizes, with fragments of shells, bryo- 
zoans, foraminiferans, old worm tubes, algae. 

Remarks. — The tj^pe of T. hamatus Moore from Alaska was ex- 
amined and is herein referred to T. cincinnatus; it is a small specimen, 
consisting of anterior end only. 

New records. — Arctic Alaska: Off Point Barrow base, up to 15 
miles from shore, 20-123.5 fms,, on bottoms of mass of worm tubes, 
rocks, stones, gravel, attached to stems of Tuhularia (6 stations, 7 
specimens). West Greenland: Godhavn, off Hare Island, 70°20' 



328 PROCEEDINGS OF THE NATIONAL MUSEUM vol.108 

N., 56° W., 90 fms., Greely Relief Expedition, 1884. One mile 
northwest Conical Rock, 25-60 fms., and north shore Wolstenholm, 
13-25 fms., Bartlett, 1940. East Greenland: Off Cape Hold with 
Hope, 23-40 fms., Bartlett, 1939. East Coast North America: 
Off Labrador, 12-30 fms.. Blue Dolphin Expedition, 1949; off New 
Brunswick, Nova Scotia, Maine, New Hampshire, Massachusetts, 16- 
640 fms., U. S. Fish Commission. West Coast North America: 
Washington and Puget Sounds, low water to 46 fms., mud, shell, rock, 
Pettibone. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Jan Mayen, Spitsbergen, Franz 
Josef Land, Barents Sea, Novaya Zemlya. Also Iceland, Faroes, 
Shetlands, Norway to Portugal, Azores, Madeira, Mediterranean, 
Adriatic; Labrador to Massachusetts; Bering Sea to Washington; 
Japan; Indian Ocean (Andamans). Southern latitudes: South Geor- 
gia, South Orkneys, South Shetlands, Graham Coast, Palmer 
Peninsula. In low water to 1,391 fathoms. 

Genus Polycirrus Grube, 1851 
Polycirrus medusa Grube, 1855 

Figure 37, e, f 

Polycirrus medusa Grube, 1855, p. 120. — Hessle, 1917, p. 220. — Fauvel, 1927, p. 
279, fig. 97, a-d.— Augener, 1928, p. 795.— Annenkova, 1934, p. 322; 1937, 
p. 194; 1938, p. 210.— Gorbunov, 1946, p. 39.— Zatsepin, 1948, p. 154, pi. 38, 
fig. 5.— Wesenberg-Lund, 1950a, p. 54; 1950b, p. 123; 1951, p. 114. 

Ereutho smitti Malmgren, 1865, p. 391, pi. 23, fig. 63. — Webster and Benedict, 
1887, p. 749. 

Polycirrus sp., Hartman, 1948, p. 45. 

Description. — Length up to 70 mm., width 6 mm. Integument 
tesselated. Large unpaired ventral shield on first few segments, 
followed by about six pairs of ventral shields separated in midline. 
Prostomium and buccal segment forming lower lip, wide semicircular 
dorsal cephalic ridge (without eye-spots) , large trilobed or undulating 
tentacular membrane with very numerous tentacles on its outer edge, 
the inner part forming folded upper lip; some tentacles long, slender, 
filiform, others thicker, distinctly grooved. Thoracic setigers usually 
13 (10-13), beginning on segment 3 (or 2). Without thoracic uncini; 
uncini begin on segment 16 (or 15 or just posterior to last setiger), 
uniserial, few in number, very small and inconspicuous, on slightly 
raised abdominal pinnules. Six pairs nephridial papillae on segments 
3-8. Females with large, yolky eggs (Point Barrow, September 15, 
1948, and October 14, 1949). Color: In life: Red. In alcohol: 
Colorless. Tube : Simple galleries in mud or sand. 

New records. — Arctic Alaska: Off Point Barrow base, up to 15 
miles from shore, 13.3-78.2 fms., on bottoms of mud and various com- 



MARINE POLYCHAETE WORMS — PETTIBONE 329 

binations of mud, pebbles, gravel, stones, rocks, large perforated 
rocks, with bryozoans, worm tubes (13 stations, 35 specimens). 
Spitsbergen: Spitsbergen Sea, U. S. S. Alliance, 1881. Northwest 
Greenland: One mile northwest Conical Rock, 25-60 fms., Bartlett, 
1940. Bering Sea: St. Paul Island, Pribilofs, Palmer, 1890. East 
Coast North America: Off Labrador, 45 fms., silt, Blue Dolphin 
Expedition, 1949. West Coast North America: Pavlof Bay, 
Alaska, 150 fms., Alaska King Crab Investigation, 1940; Washington 
Sound, 12-46 fms., mud and mussels, Pettibone. 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Spitsbergen, White Sea, Novaya Zemlya, 
Kara Sea. Also Iceland, Faroes, Swedish west coast to France, 
Mediterranean; Labrador to Maine; Bering Sea to Washington; 
Okhotsk Sea to north Japan Sea. In low water to 889 fathoms. 

Genus Trichohranchus Malmgren, 1865 
Trtchobranchus glacialis Malmgren, 1865 

Figure 37, g-i 

Trichohranchus glacialis Malmgren, 1865, p. 395, pi. 24, fig. 65. — Webster and 
Benedict, 1887, p. 750.— Ehlers, 1913, p. 566.— Hessle, 1917, p. 131.— 
Fauvel, 1927, p. 288, fig.lOO,a-h.— Augener, 1928, p. 792.— Annenkova, 
1937, p. 190; 1938, p. 202.— Zatsepin, 1948, p. 153, pi. 38, fig. 1.— Wesenberg- 
Lund, 1950a, p. 55; 1950b, p. 125; 1951, p. 115.— Berkeley and Berkeley, 
1952, p. 76, figs. 154, 155. 

Trichohranchus glacialis var. antarcticus Hessle, 1917, p. 132. — Hartman, 1952, 
p. 233. 

Description. — Length up to 30 mm., width 3 mm., segments 60-70. 
Buccal segment and prostomium form thick, tesselated, inflated lower 
lip connected laterally with pair of projecting, rounded, flattened lobes 
(ventral to mass of tentacles), cephalic ridge with numerous black 
eyespots (absent in var. antarcticus) , undulating tentacular membrane 
with very numerous tentacles, and folded upper lip; some tentacles 
larger, distinctly grooved; others smaller, filiform. Three pairs 
branchiae on segments 2-4, each composed of single long filament. 
With slightly projecting lateral lobes on branchial segments. Thoracic 
setigers and uncinigers 15, beginning on segment 6; notosetae limbate, 
with smooth capillary tips; neuropodial uncini aciculiform (crotchets). 
Abdominal segments with avicular uncini in single rows on triangular 
projecting pinnules. Pygidium with anal opening crenulate. Color- 
less in alcohol. Tube membranous, encrusted with mud or fine sand. 

New records. — Arctic Alaska: Off Point Barrow base, up to 5 
miles from shore, 27-49 fms., on various combinations of mud, gravel, 
stones, rocks, shells (5 stations, 15 specimens). East Coast North 
America: Cape Cod Bay, Massachusetts, 27-118 fms., U. S. Fish 
Commission. 



330 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Distribution. — Widely'^distributed in the Arctic: Alaskan Arctic, 
Greenland, Spitsbergen, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes, Swedish west coast to Spain, Canary Islands, Mediterranean; 
Maine to Massachusetts; British Columbia; north Japan Sea. Var. 
antarcticus: Magellan Straits, South Georgia, Ross Island, Kaiser 
Wilhehn II Land. In 3-1,517 fathoms. 

Genus Terehellides M. Sars, 1835 
Terebellides stroetnii M. Sars, 1835 

FiQUEB 37, j-m 

Terebellides stroemii M. Sars, 1835, p. 48, pi. 13, fig. 31. — Malmgren, 1865, p 
396, pi. 20, fig. 48.— Webster and Benedict, 1887, p. 750.— Moore, 1903, p 
478; 1908, p. 352; 1923, p. 199.— Hessle, 1917, p. 137.— Chamberlin, 1920 
p. 23.— Eliason, 1920, p. 72.— Fauvel, 1927, p. 291, fig. 100, i-q; 1932, p 
234; 1947, p. 79, fig. 76, f-i.— Augener, 1928, p. 797.— Gustafson, 1936, p 
9.— Annenkova, 1937, p. 190; 1938, p. 202.— Treadwell, 1937, p. 35.— Okuda, 
1938b, p. 102.— Friedrich, 1939, p. 127.— Berkeley and Berkeley, 1943, p 
130; 1944, p. 5; 1952, p. 75, figs. 152, 153.— Hartman, 1944a, pp. 336, 343 
1944d, p. 24; 1951, p. 113.— Thorson, 1946, p. 124, fig. 69.— Gorbunov, 1946 
p. 39.— Zatsepin, 1948, p. 153, pi. 38, fig. 3.— St0p-Bowitz, 1948c, p. 68.— 
Wesenberg-Lund, 1949, p. 355; 1950a, p. 55; 1950b, p. 126; 1951, p. 115, fig 
11. — Hartman and Reish, 1950, p. 44. 

Terebellides stroemii var. japonica Moore, 1903, p. 478; 1908, p. 352; 1923, p. 200 

Description. — Length up to 75 mm., width 8 mm., segments 50-60 
Prostomium and buccal segment forming large, transverse, crescent- 
shaped plate below mouth, large and undulating tentacular membrane 
with numerous, short, grooved tentacles, folded upper lip; dorsaUy 
without cephalic ridge or eye-spots. Segment 2 short, achaetous. 
Single branchia formed of large cylindrical trunk dorsally on segment 
3 (first setiger) and four pectinate lobes, frequently interlocked giving 
appearance of more or less solid mass. With somewhat developed 
lateral lobes extending ventraUy on segments 2-7 (larger on segments 
3-6). Thoracic setigers 18, begin on segment 3. Uncini begin on 
segment 8 (setiger 6), those on first unciniger consisting of crotchets 
with long manubrium and bent, pointed tips, those on rest of thoracic 
segments long, aciculiform, with large fang and few denticles; may be 
crowded in more than one row in large specimens. Abdominal 
uncini short, avicular, pectiniform, in single rows on projecting tri- 
angular pinnules. Prominent paired nephridial papillae on segment 3, 
smaller ones on segments 6 and 7. Pygidium with terminal anus, with 
sides crenulate. Female with large yolky eggs (ripe eggs and sperm, 
September 26, 1949). Color: In life: Body orange, branchia orange- 
red, tentacles tan. In alcohol: Colorless, with smooth iridescent 
surface. 

Parasites. — One of the 87 specimens had the parasitic copepod 
Saccopsis terebellidis Levinsen (identified by P. L. lUg) attached to 



MARINE POLYCHAETE WORMS — PETTIBONE 



331 




Figure 37. — Terebellidae : a, Leaena abranchiata, lateral view anterior end; b, same, notoseta; 
c, Lanassa venusta, notoseta; d, Thelepus cincinnatus, avicular uncinus; e, Polycirrus 
medusa, dorsal view anterior end;/, same, ventral view anterior end; g, Trichobranchus 
glacialis, dorsal view anterior end; h, same, thoracic aciculiform uncinus; i, same, abdom- 
inal avicular uncinus; ;', Terebellides stroemii, lateral view anterior end; k, same, crotchet 
from first unciniger; /, same, thoracic aciculiform uncinus; m, same, abdominal avicular 
uncinus. (For explanation of symbols, see p. 210.) 



261112—54- 



332 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

anterior part in the region of the branchia. The same polychaete and 
copepod species were found in Iceland waters and figured by Wesen- 
berg-Lund (1951, p. 117, fig. 11). 

New records. — Arctic Alaska: West side Elson Lagoon near Point 
Barrow, 1.2 fms,; Eluitkak Pass, Elson Lagoon; Point Barrow base, 
washed ashore; off Point Barrow base, up to 15 miles from shore, 18.3- 
123.5 fms., on bottoms of mud, stones, mass of worm tubes, and various 
combinations of mud, sand, pebbles, gravel, stones, rocks, large per- 
forated rocks, with shells, bryozoans, worm tubes (26 stations, 87 
specimens). West Coast North America: Washington and Puget 
Sounds, 10-165 fms., Pettibone. East Coast North America: Off 
Labrador, 5-65 fms.. Blue Dolphin Expeditions, 1949, 1950; off Nova 
Scotia, Maine, Massachusetts, Rhode Island, Long Island Sound, 
6-368 fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Jan Mayen, Spitsbergen, Franz 
Josef Land, Barents Sea, White Sea, Novaya Zemlya, Kara Sea. Also 
Iceland, Faroes, Norway to Portugal, Mediterranean, Adriatic, Black 
Sea; Hudson Bay to Long Island Sound, Gulf of Mexico, West Indies; 
Bering Sea to southern California, Panamd, Venezuela; north Japan 
Sea to Japan; Iranian Gulf; Indian Ocean; South Africa, Bouvet 
Island, Kerguelen, New Caledonia. In low water to 1,611 fathoms. 

Family Sabellidae 

Body cylindrical, tapered posteriorly. Prostomium indistinct. 
Buccal region with mouth terminal, with dorsal lip, two ventral lips 
or vesicular bulbs, two fleshy, membranous or filiform palps of variable 
length. Branchiae form conspicuous, often highly colored terminal 
funnellike plume surrounding mouth; plume formed of two semi- 
circular or spiral lobes bearing numerous filaments each with two rows 
of ciliated barbules (fig. 38, a). Branchiae without operculum. A 
collarette more or less developed on first segment. Body divided 
into two regions: (1) thoracic; few segments (4-12), with dorsal bun- 
dles of capillary setae and ventral uncinigerous tori (tori lacking in 
Myxicola); without thoracic membrane; (2) abdominal; with inversion 



Figure 38. — Sabellidae: a, Sabella crassicornis, lateral view anterior end; b, same, tip 
of branchial filament; c, same, thoracic notosetae; d, same, dorsal view pygidium;^, same, 
thoracic hoe-like seta from ventral torus;/, same, thoracic avicular uncinus from ventral 
torus; g, same, ventral view collarette; h, same, lateral view; i, same, dorsal view; /, Pot- 
amilla neglecta, dorsal view collarette; k, same, lateral view; /, same, ventral view; m, 
same, dorsal thoracic limbate seta; n, same, dorsal thoracic spatulate setae; o, Potamilla 
reniformis, ventral view collarette; p, same, lateral view; q, same, dorsal view; r, same, 
portion of branchial filament; s, same, thoracic avicular uncinus from ventral torus; t, 
same, thoracic hoe-like seta from ventral torus; «, same, tip of tube, rolled at free end. 
(For explanation of symbols, see p. 210.) 



MARINE POLYCHAETE WORMS — PETTIBONE 

I 



333 




Figure 38. — For explanation see facing page. 



334 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

of setae, dorsal uncinigerous tori and ventral capillary setae. A 
median ciliated faecal groove runs length of body, ventral in abdom- 
inal region and crossing to dorsal surface in thoracic region. Tube 
cylindrical, permanent or more or less transitory, gelatinous, mem- 
branous, or horny, covered or not with mud, sand, gravel, debris of 
shells. 

Represented by five genera and seven species at Point Barrow. 
All the genera have large branchial plumes composed of two similar 
semicircular lobes; branchial filaments without large subterminal 
eyes, without dorsal appendages. First setigerous segments with 
limbate notosetae only; neurosetae begin on setiger 2. Abdominal 
region with numerous segments. 

Key to the genera of SaheUidae from Point Barrow 

1. Ventral tori of thorax with row of uncini of the avicular form (fig. 38, /; in 

Sabella and Potamilla with additional row of hoelike setae, fig. 38, t) . Col- 
larette well developed. Branchial filaments with or without eyes, united 

at base only, with palmar membrane poorly developed or lacking 2 

Ventral tori of thorax with crotchets with long manubrium (fig. 39, t; without 
tori in Myxicola). Branchial filaments without eyes, united along a large 
part of their length by a well-developed palmar membrane (fig. 39, w) — 3 

2. Dorsal thoracic setae all limbate, some long with straight borders, others short 

with wide borders (fig. 38, c) Sabella (p. 334) 

Dorsal thoracic setae of two kinds, some limbate, others spatulate (fig. 38, m, n) 

Potamilla (p. 335) 

3. Uncinigerous tori projecting, with short, lateral rows of uncini. Collarette 

well developed (fig. 39, a-c). Dorsal thoracic setae of two kinds, limbate 

and spatulate (fig. 39, /, g) 4 

Without projecting uncinigerous tori, abdominal uncini in almost complete 
transverse band (fig. 39, p). Collarette poorly developed, represented by 
a triangular ventral lobe (fig. 39, o). Dorsal thoracic setae all capillary, 
limbate Myxicola (p. 340) 

4. Posterior segments without ventral depression Chone (p. 337) 

Posterior segments about 10, with ventral suckerlike disc, large ventral de- 
pression with flared sides (fig. 39, n) Euchoiie (p. 339) 

Genus Sabella Linn^, 1767 (sensu Malmgren, 1865) 

Sabella crassicornis Sars, 18.51 

Figure 38, a-i 

Sabella crassicornis Sars, 1851, p. 202.— Malmgren, 1865, p. 399, pi. 27, fig. 83.— 

Moore, 1909b, p. 144.— Johansson, 1927, p. 119.— Hartman, 1942a, p. 78; 

1948, p. 46.— Berkeley and Berkeley, 1943, p. 130; 1952, p. 114, figs. 236, 

237.— Zatsepiu, 1948, p. 161, pi. 39, fig. 4. 
Sabella fabricii Kroyer, 1856, p. 20.— Fauvel, 1927, p. 300, fig. 103, a-g.— Augener, 

1928, p. 800.— Annenkova, 1934, p. 322; 1937, p. 195; 1938, p. 211.— Wesen- 

berg-Lund, 1950b, p. 128; 1951, p. 118. 
Sabella spetsbergensis Malmgren, 1865, p. 399, pi. 29, fig. 93. 
Sabella spitzbergensis Webster and Benedict, 1887, p. 750. 



MARINE POLYCHAETE WORMS — PETTIBONE 335 

Description. — Length 20-80 mm., width 3-4 mm. Collarette 
widely separated middorsally, with midventral slit and lateral notches, 
resulting in 4-lobed structm-e; ventral lobes closely approximated, 
may be deflected. Two branchial lobes each with about 16 filaments 
(15-35), with short tapering tips. Branchial filaments with usually 
four to six (2-8) pairs of eyes (located in color bands of filaments and 
may easily be overlooked). Thoracic setigers usually 8 (7-9). Abdo- 
men with one or two pairs of eye-spots between parapodial rami, well 
developed toward posterior end. Pj^gidium with pair of bulbous 
lobes with groups of eye-spots on dorsolateral sides. Color: In 
life : Body flesh color, with branchial filaments banded rusty red, 4-7 
transverse bands per filament. In alcohol : Body colorless with red- 
dish purple bands on branchial filaments. Tube: Cylindrical, free 
end flexible, covered with thin, smooth layer of mud; embedded part 
transparent, horny, rigid, covered with sand grains and foreign 
material. 

New records. — Arctic Alaska: Off Point Barrow base, up to 15 
miles from shore, 36-123.5 fms., on bottoms of stones, mass of worm 
tubes, and various combinations of gravel, stones, rocks, large per- 
forated rocks, with worm tubes (8 stations, 15 specimens). Kam- 
chatka: Petropavlovsk, Grevnitzky, 1888. Alaska: New Harbor, 
Unga Island, under stones, Dall, 1872; Albatross Sta. 2847, 55°01' N., 
160°18' W., 48 fms., and station at Kodiak, 1888. Canadian Arctic: 
Foxe Basin, 25-31 fms., Bartlett, 1927. East Coast North America: 
Off Labrador, 5-6 fms.. Blue Dolphin Expedition, 1949; Bay of Fundy, 
Maine, Massachusetts, 15 fms., U. S. Fish Commission. Central 
America: Golfo Dulce, west Costa Rica, M. Valerio. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Novaya Zemlya, Kara 
Sea. Also Iceland, Faroes, Norway to France, Mediterranean; 
Hudson Bay to Massachusetts; Bering Sea to California, Central 
America (Costa Rica); north Japan Sea to Japan. In low water to 
230 fathoms. 

Genus Potaniilla Malmgren, 1865 

Key to the species of Potamilla from Point Barrow 

1. Without branchial eyes P. neglecta 

With compound eyes in single rows on branchial filaments, 0-8 per filament 
(fig. 38, r) P. reniformis 

Potamilla neglecta (Sars, 1851) 

Figure 38, j-n 

Sabella neglecta Sars, 1851, p. 203. 

Potamilla neglecta Malmgren, 1865, p. 401, pi. 27, fig. 84. — Webster and Benedict, 
1884, p. 736.— Moore, 1909b, p. 145; 1923, p. 242.— Johansson, 1927, p. 143.— 



336 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Augener, 1928, p. 801.— Hartman, 1942a, p. 81; 1944a, pp. 336, 343; 

1948, p. 46.— Zatsepin, 1948, p. 162, pi. 39, fig. 7.— Wesenberg-Lund, 1950a, 

p. 56; 1950b, p. 128; 1951, p. 119.— Berkeley and Berkeley, 1952, p. 116, fig. 

238. 
Potamilla torelli Malmgren, 1865, p. 402; 1867, p. 114, pi. 13, fig. 76.— Ehlers, 

1913, p. 575.— Fauvel, 1927, p. 310, fig. 107, m-s; 1934a, p. 71.— Annenkova, 

1934, p. 322; 1937, p. 195; 1938, p. 213.— Wesenberg-Lund, 1951, p. 121. 
Potamilla acuminata Moore and Bush, 1904, p. 159, pi. 11, figs. 3-6; pi. 12, fig. 

41. 

Description. — Length 30-84 mm., 1-3 mm. wide. Collarette widely- 
separated dorsally, sloping ventrally, with midventral slit, entire 
laterally, resulting in a 2-lobed structure; ventral lobes triangular, 
may be deflected. Branchial lobes each with about 16 filaments 
(6-20), with short, naked distal tips, without eyes. Thoracic setigers 
usually 8 (5-8). Pygidium with pair of bulbous lobes with dark eye- 
spots. Color: In alcohol: Body colorless, branchiae with two or 
three difiFused reddish brown bands. Tube: Horny, transparent, 
more or less covered with sand or mud; smaller tubes with rather 
uniform layer of small sand grains; larger tubes may be encrusted 
with living barnacles, bryozoans, hydroids, foraminiferans, amphipod 
tubes. Large yolky eggs in a single layer, pressed close to wall, 
about one-third way down in tube; a transparent, thin membrane 
laid down between eggs and worm (large developing yellow eggs 
found in tube, September 6, 1949, dredged in 36 fathoms. Point 
Barrow) . 

New records. — Arctic Alaska: Off Point Barrow base, up to 12.1 
miles from shore, 21.7-123.5 fms., on bottoms of mass of worm tubes 
and pebbles, gravel, rocks, large stones (7 stations, 24 specimens). 
Bering Sea: Albatross Sta. 3496, 56°32' N., 169°45' W., 41 fms., 
1893; St. Paul Island, Pribilofs, Hahn, 1911. Alaska: Albatross 
Sta., Kodiak, 1888. West Greenland: Off Hare Island, 70°20' N., 
56° W., 90 fms., U. S. S. Alert, 1884. East Coast North America: 
Off Labrador, 125 fms., Blue Dolphin Expedition, 1949; off Maine, 
Massachusetts, 31-130 fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Spitsbergen. Also Iceland, Faroes, Nor- 
way to France, Madeira, Mediterranean, Adriatic, Cape Verde 
Islands; Labrador to Massachusetts; Bering Sea to California; north 
Japan Sea to Japan; Antarctic. In low water to 1,044 fathoms. 

Potamilla reniformis (Leuckart, 1849) 

Figure 38, o-u 

Sabella reniformis Leuckart, 1849, p. 183, pi. 3, fig. 8. 

Potamilla reniformis Malmgren, 1867, p. 114, pi. 13, fig. 77. — Webster and Bene- 
dict, 1884, p. 736; 1887, p. 750.— Johansson, 1927, p. 142.— Fauvel, 1927, 



MARINE POLYCHAETE WORMS — PETTIBONE 337 

p. 309, fig. 107, a-1.— Okuda, 1937b, p. 61.— Annenkova, 1934, p. 322; 1937, 

p. 195; 1938, p. 212.— Zatsepin, 1948, p. 161, pi. 39, fig. 6.— Wesenberg-Lund, 

1950a, p. 57; 1950b, p. 129; 1951, p. 120. 
Pseudopoiamilla intermedia Moore, 1905c, p. 562, pi. 37, figs. 15-22; 1908, p. 359. — 

Hartman, 1938c, p. 25, pi. 2, fig. 8; pi. 3, figs. 1-4; 1948, p. 46.— Rioja, 

1941, p. 732. 
Pseudopotamilla reniformis Hartman, 1944a, pp. 336, 343, pi. 21, figs. 3, 6; 1945, 

p. 47.— Berkeley and Berkeley, 1952, p. 116, fig. 239. 

Description. — Length 80-100 mm., width 1-2 mm. Collarette 
with middorsal depression, deeply notched dorsolaterally (at level of 
notopodia), with midventral slit, resulting in a 4-lobed structm-e of 
pair of rounded or triangular dorsal lobes and pair of rounded ventral 
lobes which may be deflected. Branchial lobes each with about 10 
filaments (5-15). At least some of branchial filaments with large 
compound eyes, usually 0-3 per filament (0-8), eyes may be variable 
in size or subequal, deep reddish in alcohol. Thoracic setigers usually 
10 (7-14). Pygidium with pair of bulbous lobes with eye-spots. 
Color: In alcohol: Body colorless, branchiae with diffused reddish 
brown bands in region of eyes. Tube: Somewhat twisted, horny, 
translucent or opaque, more or less covered with sand or mud, the 
free end usually flattened, rolled scroll-like when animal pulls inside. 

New records. — Arctic Alaska: Off Point Barrow base, up to 7.4 
miles from shore, 21.7-54.6 fms., on bottoms of pebbles, gravel, 
rocks, in holes of large stones (2 stations, 15 specimens). Bering 
Sea: Anchorage, Cape Etoliu, Nunivak Island, 8 fms., Dall, 1874. 
Southeastern Alaska: Lituya Bay, 6-9 fms., Dall, 1874. East 
Coast North America: Off Nova Scotia, Maine, New Hampshire, 
Massachusetts, Rhode Island, Delaware, 10-317 fms., U. S. Fish 
Commission. 

Distribution. — Scattered records in the Arctic: Arctic Alaska, 
Greenland. Also Iceland, Swedish west coast to France, Mediter- 
ranean; Nova Scotia to North Carolina; Bering Sea to British Co- 
lumbia, Mexico; north Japan Sea to Japan. In low water to 317 
fathoms. 

Genus Chone Kroyer, 1856 

Both species have the segments biannulate. Collarette deeply 
incised middorsally, forming two longitudinal-folded brackets, with 
sides smooth; without midventral or lateral notches (fig. 39, a-c). 
Branchial filaments united on great part of their length by palmar 
membrane (fig. 39, m). Thoracic setigers 8. Thoracic dorsal setae 
of three kinds — limbate capillary setae, fine bayonette setae, and 
spatulate setae with asymmetrical mucronate tips or gently rounded 
(fig. 39,f-h). Pygidium with anus terminal, with rounded to conical 
bulbous dorsal valve (fig. 39, d). 



338 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Key to the species of Chone from Point Barrow 

1. Collarette nearly straight (fig. 39, b). Free end of branchial filaments flat- 
tened, foliaceous, widely limbate (fig. 39, e)_- -_C. infuudibuliformis 

Collarette oblique, longer on ventral side (fig. 39, k). Free end of branchial 
filaments with long filiform tips (fig. 39, I) C. duneri 

Chone infundibuliformia Kroyer, 1856 

Figure 39, a-j 

Chone infundibuliformis Kroyer, 1856, p. 33. — Malmgren, 1865, p. 404, pi. 28, 
fig. 87.— Th6el, 1879, p. 66.— Wir6n, 1883, p. 422.— Fauvel, 1927, p. 334, 
fig. 116, a-o.— Augener, 1928, p. 806.— Annenkova, 1934, p. 322; 1937, 
p. 196; 1938, p. 215.— Hartman, 1942b, p. 136; 1944a, pp. 334, 336 (not pi. 
20, fig. 5), pi. 21, fig. 7.— Berkeley and Berkeley, 1943, p. 130; 1952, p. 123, 
figs. 252, 253.— Gorbunov, 1946, p. 39.— Zatsepin, 1948, p. 164, pi. 39, 
fig. 13.— Wesenberg-Lund, 1950a, p. 58; 1950b, p. 131; 1951, p. 123. 

Chone gracilis Moore, 1906b, p. 257, pi. 12, figs. 62-66. — Berkeley and Berkeley, 
1952, p. 123, fig. 254. 

Description. — Length 30 mm., width 2.5 mm. (up to 120 mm. long, 
6 mm. wide — Fauvel, 1927). Collarette nearly straight laterally. 
Branchial filaments about 15 (10-36), with tips more or less foliaceous, 
edged by transparent border (prolongations of palmar membrane). 
Color: In life: Variable — flesh color, branchiae orange spotted with 
white; olive green with distal half of branchiae brick red, basal half 
red and olive, chalk white on outer sides of bases of branchiae. In 
alcohol: Colorless, gra3dsh, or tannish. Tube: Membranous, en- 
crusted with sand, mud, or pebbles of variable sizes, foraminiferans. 

New records. — Arctic Alaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, up to 7.5 miles from 
shore, 20^9 fms., on bottoms of mud, stones, and various combina- 
tions of mud, sand, gravel, stones, rocks, large perforated rocks, shells 
(12 stations, 32 specimens). Canadian Arctic: East side Cobourg 
Island, Baffin Bay, 75° 40' N., 78° 40' W., Bartlett, 1935. North- 
west Greenland: 1 mile northwest Conical Rock, Bartlett, 1940. 
East Greenland: Off Cape Hold with Hope, 23-40 fms., Bartlett, 
1939. Spitsbergen: Spitsbergen Sea, U.S.S. ^Z/iance, 1881. Bering 
Sea: St. Paul Island, Pribilofs, Palmer, 1890; St. George Island, 
Pribilofs, Hanna, 1913; Albatross Sta. 3232, 58° 30' N., 157° 34' W., 
10.5 fms., 1890, Sta. 3233, 58° 23' N., 157° 42' W., 7 fms., 1890, and 
Sta. 3289, 56° 44' N., 159° 16' W., 16 fms., 1890; Kiska, Aleutians, 
Dall. West Coast North America: Strait of Juan de Fuca, Wash- 
ington, 40 fms., mud, Pettibone. East Coast North America: Off 
Nova Scotia, Maine, Massachusetts, Rhode Island, 10-206 fms., U. 
S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Novaya Zemlya, Kara 



MARINE POLYCHAETE WORMS — PETTIBONE 339 

Sea. Also Iceland, Faroes, Shetlands, Scandinavian coast to Danish 
waters; Hudson Bay to Rhode Island; Bering Sea to California; north 
Japan Sea. In low water to 1,955 fathoms. 

Chone dun^ri Malmgren, 1867 

Figure 39, k, I 

Chone dunSri Malmgren, 1867, p. 116, pi. 13, fig. 75.— Wir^n, 1883, p. 422.— 
Fauvel, 1927, p. 336, fig. 117, 1-r; 1934a, p. 74.— Augener, 1928, p. 807.— 
Monro, 1936, p. 189.— Treadwell, 1937, p. 35.— Berkeley and Berkeley, 1944, 
p. 5. — Zatsepin, 1948, p. 164, pi. 39, fig. 15. — Wesenberg-Lund, 1950a, p. 
58; 1950b, p. 130; 1951, p. 123.— Hartman, 1951, p. 117. 

Description. — Length 20-35 mm., width 1.5-2.5 mm. Collarette 
oblique, longer on ventral side. Branchial filaments about 10 (5-11), 
with long, tapered tips. Color: In life: Branchiae yellow. In alco- 
hol: Colorless or tannish. Tube: Thin, membranous, encrusted with 
sand. 

New records. — ^ Arctic Alaska: Off Point Barrow base, up to 7.5 
miles from shore, 13.3-49 fms., on bottoms of mud and various com- 
binations of mud, sand, gravel, stones, rocks, large perforated rocks, 
shells (9 stations, 21 specimens). Canadian Arctic: Baffin Island, 
66° 43' N., 80° 07' W., Bartlett, 1927. East Coast North America: 
Off Labrador, 8 fms., mud. Blue Dolphin Expedition, 1949. 

Distribution. — Widely distributed in the Arctic: Alaskan and Ca- 
nadian Arctic, Greenland, Spitsbergen, Novaya Zemlya, Kara Sea. 
Also Iceland, Faroes, Norway to North Sea, Madeira, Mediterranean, 
Adriatic; Labrador; Florida; west coast South America (Perii). In 
8-889 fathoms. 

Genus Euchone Malmgren, 1865 
Euchone analis (Kroyer, 1856) 

Figure 39, m, n 

Sabella analis KrSyer, 1856, p. 17. 

Euchone analis Malmgren, 1865, p. 406, pi. 28, fig. 88.— Th6el, 1879, p. 65.— 
Wir6n, 1883, p. 423.— Chamberlin, 1920, p. 27.— Augener, 1928, p. 804.— 
Annenkova, 1937, p. 196; 1938, p. 216.— Berkeley and Berkeley, 1943, p. 
130; 1952, p. 121, figs. 250, 251.— Gorbunov, 1946, p. 39.— Zatsepin, 1948, 
p. 163, pi. 39, fig. 12.— Wesenberg-Lund, 1950a, p. 59; 1950b, p. 132; 1951, 
p. 124. 

Description. — Length 19-28 mm., width 1.8 mm. (up to 60 mm. 
long — Augener, 1928). Segments biaimulate and, with the ventral 
faecal groove, forming four large areas per segment. Collarette 
with opening dorsally, nearly straight and entire laterally, with 
shallow midventral incision. Branchial filaments about 10 per lobe 
(up to 14), united through a great part of their length by a well- 
developed palmar membrane. Thoracic setigers 8. Ventral anal 



340 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i03 

depression extending over 10-12 most posterior segments, bordered 
by high sinuous membrane. Pygidium with terminal anus, with 
conical bulbous dorsal valve. Colorless or tannish in alcohol. Tube 
membranous, encrusted with sand grains and certain amount of 
debris. 

New records. — Arctic Alaska: Off Point Barrow base, up to 12.1 
miles from shore, 36-123.5 fms., on bottoms of mass of worm tubes 
and stones, rocks, large perforated rocks (3 stations, 3 specimens). 
East Coast North America: Off Labrador, 30 fms., mud with 
rock, Blue Dolphin Expedition, 1950. 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Jan Mayen, Spits- 
bergen, Novaya Zemlya, Kara Sea. Also Iceland, Norway to 
Danish waters; Hudson Bay to Labrador; Bering Sea to British 
Columbia; north Japan Sea. In 2-389 fathoms. 

Genus Myxicola (Koch in MS.) Renier, 1848 
Myxicola infundibulum (Montagu, 1808) 

Figure 39, o-q 

Amphitrite infundibulum Montagu, 1808, p. 109, pi. 8. 

Myxicola steenstrupii Kroyer, 1856, p. 35. — Malmgren, 1865, p. 408, pi. 29, fig. 

90.— Th^el, 1879, p. 66.— Webster and Benedict, 1884, p. 737; 1887, p. 

750.— Moore, 1909b, p. 144.— Eliason, 1920, p. 79.— Zatsepin, 1948, p. 164, 

pi. 39, fig. 16. 
Myxicola infundibulum Fauvel, 1927, p. 342, fig. 119, a-i; 1934a, p. 74; 1936, 

p. 86.— Annenkova, 1934, p. 322.— Okuda, 1939, p. 243.— Hartman, 1944a, 

pp. 335, 343, pi. 21, fig. 1; 1948, p. 47.— Wesenberg-Lund, 1950b, p. 

134; 1951, p. 125.— Berkeley and Berkeley, 1952, p. 119, fig. 244. 

Description. — Length 23 mm., width 5 mm. (up to 200 mm. long, 
15 mm. wide — Fauvel, 1927). Collarette represented by a triangular 
ventral lobe extending anteriorly between the branchial lobes, absent 
laterally and dorsally. A middorsal groove on first 8-10 segments. 
Branchial filaments 20-40, united for greater part of their length by 



Figure 39. — Sabellidae: a, Chone infundibulif omits, ventral view collarette; b, same, lateral 
view; c, same, dorsal view; d, same, dorsal view pygidium; e, same, tip of foliaceous bran- 
chial filament;/, same, thoracic dorsal limbate capillary seta; g, same, thoracic dorsal spatu- 
late seta; k, same, thoracic dorsal bayonette seta; i, same, thoracic crotchet from ventral 
torus; y, same, abdominal uncinus; k, Chone duneri, lateral view collarette; /, same, tip 
of branchial filament; m, Euchone analis, lateral view anterior end; n, same, ventral view 
posterior end; o, Myxicola infundibulum, lateral view anterior end; p, same, lateral view 
segment from abdominal region; ^, same, dorsal view pygidium. Serpulidae: r, Sptrorbis 
granulatus, dorsal view tube; s, same, operculum (after Fauvel); /, same, collar seta (seta 
of first setiger); u, Spirorbis spirillum, dorsal view anterior end extended from tube (var. 
ascendens, after Emerton); f, same, discoidal form of tube; w, same, operculum; at, same, 
collar seta (seta of first setiger, after Fauvel). (For explanation of symbols, see p. 210.) 



MARINE POLYCHAETE WORMS — PETTIBONE 



341 




V X 

FlGTJRE 39. — For explanation see facing page. 



342 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

well-developed palmar membrane; tips of filaments limbate. Thora- 
cic setigers 8 (7-9). Thoracic notopodia with limbate capillary setae; 
nem-opodia with crotchets with long manubrium (may disappear). 
Abdomen without projecting tori; avicular uncini in an almost com- 
plete circle; with capillary neurosetae. With one to several lateral 
eye-spots behind each parapodia. Pygidium with anus terminal, 
with rounded, bulbous dorsal valve with lateral groups of eye-spots. 
Color: In alcohol: Body irregularly speckled reddish brown, with 
branchiae violet distally. Tube: Gelatinous, transparent, very 
firm and elastic, may be quite thick. 

New records. — Arctic Alaska: Off Point Barrow base, up to 4,25 
miles from shore, 25-35.5 fms., on bottoms of mud with gravel (2 
stations, 2 specimens). West Coast North America: Washington 
and Puget Sounds, low water to 165 fms., Pettibone. East Coast 
North America: Off Nova Scotia, Maine, Massachusetts, 9-90 
fms., U. S. Fish Commission. 

Distribution. — Scattered records in the Arctic: Arctic Alaska, 
West Greenland, Novaya Zemlya. Also Iceland, Finland, to English 
Channel, Mediterranean, Adriatic; Nova Scotia to Massachusetts; 
Bering Sea to California; Japan. In low water to 287 fathoms. 

Family Serpulidae 

Body cylindrical or fusiform, slightly flattened. Prostomium indis- 
tinct. Buccal region with mouth terminal, with two transverse lips, 
palps absent or slightly developed. Branchiae form a terminal funnel- 
like plume around mouth, formed of two semicircular or spiralled 
lobes bearing few to numerous filaments, each with two rows of ciliated 
barbules (fig. 39, u). Usually with one or two opercula (calcareous, 
horny, or membranous) formed on the first dorsal filaments of the 
branchial lobes. Usually with a collarette on first segment. Body 
divided into two regions: (1) thoracic; with few segments (3-7), with 
dorsal bundles of capillary setae and ventral uncinigerous tori; usually 
with a thoracic membrane; (2) abdominal; with inversion of setae, 
dorsal uncinigerous tori and ventral capillary setae. A wide, shallow, 
longitudinal ciliated faecal groove, ventral in abdominal region, dorsal 
in thoracic. Tube cylindrical or polygonal, calcareous, opaque or 
rarely transparent, fixed to substratum (rarely free). 

Represented by a single genus and two species (two subgenera) at 
Point Barrow. 

C^nus Spirorbis Daudin, 1800 

Both species are of smaU size, asymmetrical and coiled. Branchial 
lobes each with 3-5 filaments. A single, partly calcareous operculum 



MARINE POLYCHAETE WORMS — PETTIBONE 343 

on smooth peduncle. Collarette large, widely separated dorsally, 
entire laterally and ventraUy. Three thoracic setigers, the first 
setiger with only dorsal setae (collar setae) and the next two with 
ventral tori in addition to dorsal setae. With well-developed thoracic 
membrane. A long achaetous region between thorax and abdomen 
(may be crowded with ova; spermatozoa develop in more posterior 
segments ; hermaphroditic) . About 20 abdominal setigers. Pygidium 
with two rounded lobes. Colorless in alcohol. Tube small (1-3 mm. 
in diameter), closely coiled, white, opaque, fixed to hydroids, bryozoa, 
algae, carapaces of crustaceans, spines of tunicates, stones, etc. 

Key to the species of Spirorbis from Point Barrow 

1. Tube with sinistral (counterclockwise) spiral coiling (with mouth of tube 
facing observer, the opening is on the left; fig. 39, r). Eggs incubated in 
operculum. Collar setae with well-developed basal crenulate wings (fig. 
39, t) S. (Laeospira) granulatus 

Tube with dextral (clockwise) spiral coiling (fig. 39, v). Eggs incubated in 
tube. Collar setae without crenulate wings (fig. 39, x) . 

S. (Dexiospira) spirillum 

Spirorbis (Laeospira) granulatus (Linn^, 1767) 

Figure 39, r-t 

Serpula granulata Linn^, 1767, p. 1266. 

Spirorbis quadrangular is Stimpson, 1854, p. 29. — Moore, 1908, p. 362. — Hartman, 
1944a, pp. 336, 343. 

Spirorbis (Laeospira) granulatus Borg, 1917, p. 28, figs. 14-16. — Fauvel, 1927, p. 
403, fig. 137, q-u.— Augener, 1928, p. 815.— Annenkova, 1937, p. 198; 1938, 
p. 219.— Berkeley and Berkeley, 1943, p. 130; 1952, p. 137, figs. 286, 287.— 
Gorbunov, 1946, p. 39.— ? Thorson, 1946, p. 139, fig. 80.— Zatsepin, 1948, 
p. 166.— Wesenberg-Lund, 1950a, p. 63; 1950b, p. 141; 1951, p. 135. 

Description. — Tube 1-2 mm. in diameter, dull chalky white, opaque, 
somewhat rugose and variable with two longitudinal ridges, one on 
each side (the opening almost quadrangular), or with three more or 
less distinct longitudinal keels, or without crests; the tube is coiled 
sinistrally, up to two coils, flatly on smooth surfaces or somewhat 
open when on rough surfaces; it may encircle a strand of hydroid and 
the free end may extend upward. Operculum with terminal plate 
strongly convex, calcareous, with long, cylindrical projection (short 
on one side, longer on opposite side), with large brood pouch for 
incubating eggs (eggs in operculum. Point Barrow, August 17, 1949, 
and February 18, 1950). Collar setae with blades finely serrated, 
with well-developed crenulate wing at base. 

New records. — Arctic Alaska: Off Point Barrow base, up to 12.1 
miles from shore, 27-123.5 fms., on bottoms of mass of worm tubes. 



344 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

and various combinations of mud, gravel, rocks, stones (3 stations, 7 
specimens). Southwestern Alaska: Kiska Harbor, Aleutians, 10 
fms., Dall, 1873; Canoe Bay, 25 fms., on old clam shell, Alaska King 
Crab Investigation, 1940. East Coast North America: Off Lab- 
rador, 30-35 fms., mud, rocks. Blue Dolphin Expedition, 1950; off 
Nova Scotia, Maine, Massachusetts, North Carolina (Cape Hatteras), 
17-34 fms., U. S. Fish Commission. 

Distribution. — Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Jan Mayen, Spitsbergen, Franz Josef 
Land, Novaya Zemlya, Kara Sea. Also Iceland, Faroes, Norway to 
France; Hudson Bay to North Carolina; Alaska to British Columbia; 
north Japan Sea. In low water to 239 fathoms. 

Spirorbis (Dexiospird) spirillum (Linne, 1758) 

Figure 39, u-x 

Serpula spirillum Linn6, 1758, p. 785. 

Spirorbis (Dexiospira) spirillum Borg, 1917, p. 20, figs. 3, 4. — Fauvel, 1927, p. 

392, fig. 132, f-p.— Augener, 1928, p. 814.— Annenkova, 1934, p. 322; 1937, 

p. 197; 1938, p. 218.— Treadwell, 1937, p. 35.— Hartman, 1942a, p. 91; 

1944a, pp. 336, 343, pi. 22, fig. 2.— Berkeley and Berkeley, 1943, p. 130; 

1952, p. 133, figs. 272-274.— Thorson, 1946, p. 138.— Gorbunov, 1946, p. 

39.— Zatsepin, 1948, p. 166.— Wesenberg-Lund, 1950a, p. 62; 1950b, p. 138; 

1951, p. 132. 
Circeis spirillum Chamberlin, 1920, p. 28. 
Dexiospira spirillum Hartraan, 1944b, p. 287; 1948, p. 51; 1951, p. 121. — Hartman 

and Reish, 1950, p. 47. 

Description. — Tube 1-3 mm. in diameter, white, opaque (may be 
somewhat translucent) , shiny, smooth, porcellaneous, coded dextrally, 
up to 3K coils; tube may be coiled flatly (discoidal form) when 
attached to smooth surfaces, or partly unrolled, the last coU or part 
of last coil raised from substratum (var. ascendens Levinsen, var. 
lucidus Morch) when attached to rough surfaces, as branching 
bryozoan colonies. Operculum with shallow, calcareous, concave, 
terminal plate, with slight projection (talon) on under side. Collar 
setae with serrate blades, without basal crenulate wings. Embryos 
incubated in tube (eggs in tube, Point Barrow, September 9, 1948, 
and August 30, 1949). 

New records. — Arctic Alaska: Off Point Barrow base, up to 8 
miles from shore, 21-75.5 fms., on bottoms of stones and various 
combinations of gravel, rocks, on bryozoa, on Hyas coarctatus, on 
spines of Boltenia echinata (8 stations, 39 specimens). Gulf of 
Alaska: Albatross Sta., Observation Island, Cordova, 1914. West 
Greenland: Godhavn, Greely Relief Expedition, 1884. East Coast 
North America: Off Labrador, 10-40 fms.. Blue Dolphin Expeditions, 
1949, 1950, 1951; off Nova Scotia, Maine, Massachusetts, Long 
Island Sound, 16-83 fms., U. S. Fish Commission. 



MARINE POLYCHAETE WORMS — PETTIBONE 345 

Distribution. — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Franz Josef Land, 
Kara Sea. Also Iceland, Faroes, Norway to France; Hudson Bay to 
Long Island Sound, southern Texas; Bering Sea to Mexico; north 
Japan Sea to Japan. In low water to 183 fathoms. 



346 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

References 

Alaejos y Sanz, Lufs. 

1905. Estudio descriptivo de algunas especies de polinoinos de las costaa 
de Santander. Mem. Soc. Espafiola Hist. Nat., vol. 3, pp. 1-76 
12 pis. 
Andrews, E. A. 

1891. Report upon the Annelida Polychaeta of Beaufort, North Carolina. 
Proc. U. S. Nat. Mus., vol. 14, pp. 277-302, pis. 12-18. 
Annenkova, N. p. 

1929. Beitrage zur Kenntnis der Polychaeten-Fauna der U.S.S.R. 1. Fam. 
Pectinariidae Quatrefages und Ampharetidae Malmgren, Ann. 
Mus. Zool. Acad. Leningrad, vol. 30, pp. 477-502, pis. 37-39. 

1931. Die Polychaeten in den Sammlungen der Jakutischen Expedition der 

Akademie der Wissenschaften der U.S.S.R. Zool. Anz., vol. 95, 
pp. 203-205, 4 figs. 

1932. Zur Fauna der Polychaeten des Leptev-Meeres. Issled. Morel 

USSR, Gosud, Gidrol. Inst., Leningrad, fasc. 15, pp. 133-139, figs. 
1-7. 
1934. Kurze (Ibersicht der Polychaeten der Litoralzone der Bering-Insel 
(Kommandor-Inseln) , nebst Beschreibung neuer Arten. Zool. 
Anz., vol. 106, pp. 322-331, 11 figs. 

1937. The polychaete fauna of the northern part of the Japan Sea. Issled. 

Morel USSR, Gosud. Gidrol. Inst., Leningrad, fasc. 23, pp. 139- 
216, 12 figs., 5 pis. 

1938. Polychaeta of the North Japan Sea and their horizontal and vertical 

distribution. In Reports of the Japan Sea Hydrobiological Ex- 
pedition of the Academy of Sciences of the USSR in 1934, pt. 1, 

pp. 81-230, 16 figs. 
1946. New species of Polychaeta from the Arctic Ocean. Trudy Drei- 

fufushchei ekspeditsii Glavsevmorputi na ledokolnom parokhode 

G. sedov 1937-1940, vol. 3, pp. 185-188, 3 figs. (Leningrad 

Vsesoiuznyl arkticheskil institut). 
Arwidsson, Ivar 

1898. Studien tiber die Familien Glyceridae und Goniadidae. Bergens 

Mus. Aarb., No. 11, pp. 1-69, 4 pis. 
1907. Studien iiber die skandinavischen und arktischen Maldaniden nebst 

Zusammenstellung der iibrigen bisher bekannten Arten dieser 

Familie. Zool. Jahrb., suppl. 9, pt. 1, pp. 1-308, 12 pis. 
1911a. Die Maldaniden. Wissenschaftlichen Ergebnisse der Schwedischen 

Sttdpolar-Expedition 1901-1903. Vol. 6, pt. 6, pp. 1-44, 2 pis. 
1911b. On some Irish Maldanidae. Proc. Roy. Irish Acad. Dublin, vol. 

29B, No. 6, pp. 209-228, pis. 17-19. 
1922. Systematic notes on some Maldanids. Kongl. Svenska Vetensk. 

Akad. Handl., vol. 63, No. 7, pp. 1-46, 2 pis. 
AsHwoRTH, James Hartley 

1910. The annelids of the family ArenicoUdae of North and South America, 

including an account of Arenicola glacialis Murdoch. Proc. U. S. 

Nat. Mus., vol. 39, No. 1772, pp. 1-32, figs. 1-14. 
1912. Catalogue of the Chaetopoda in the British Museum (Natural 

History). A. Polychaeta: Pt. I. Arenicolidae. Pp. 1-175, 68 figs., 

15 pis. 



MARINE POLYCHAETE WORMS — PETTIBONE 347 

1924. The rediscovery of Arenicola glacialis Murdoch, with observations 
on its characters and on its status. Rep. Canadian Arctic Exped. 
1913-18. Pt. J: Polychaeta, suppl. 1, vol. 9, pp. Sj-Tj, 1 fig. 
AuGENBR, Hermann 

1913. Polychaeten von Franz-Joseph-Land. I, II. Zool. Anz., vol. 41, 

pp. 202-220, 253-273. 
1928. Die Polychaten von Spitzbergen. In Fauna Arctica. Romer und 

Schaudinn, vol. 5, pt. 3, pp. 649-834, pi. 11. 
1939. Beitrag zur Polychaetenfauna der Ostsee. Kieler Meersf., vol. 3, 
pt. 1, pp. 133-147, 3 figs. 
Bergstrom, Erik 

1914. Zur Systematik der Polychaetenfamilie der Phyllodociden. Zool. 

Bidr. Uppsala, vol. 3, pp. 37-224, 81 figs., pis. 1-5. 
Berkeley, Edith 

1930. Polychaetous annelids from the Nanaimo District. 5. Ammocha- 

ridae to Myzostomidae. Contr. Canadian Biol. Fish., new ser., 

vol. 6, No. 5, pp. 67-77, fig. 1. 
Berkeley, Edith, and Berkeley, Cyril 

1936. Notes on Polychaeta from the coast of western Canada. I. Spionidae. 

Ann. Mag. Nat. Hist., ser. 10, vol. 18, pp. 468-477, 1 fig. 
1938. Notes on Polychaeta from the coast of western Canada. II. Syllidae. 

Ann. Mag. Nat. Hist., ser. 11, vol. 1, pp. 33-49, 12 figs. 

1942. North Pacific Polychaeta, chiefly from the west coast of Vancouver 

Island, Alaska, and Bering Sea. Canadian Journ. Res., vol. 20, 
pp. 183-208, 6 figs. 

1943. Biological and oceanographical conditions in Hudson Bay. II. 

Polychaeta from Hudson Bay. Journ. Fish. Res. Board Canada, 
vol. 6, No. 2, pp. 129-132. 

1944. Polychaeta from the western Canadian Arctic region. Canadian 

Journ. Res., vol. 22, pp. 1-5, 6 figs. 

1945. Notes on Polychaeta from the coast of western Canada. III. Ann. 

Mag. Nat. Hist., ser. 11, vol. 12, pp. 316-335, 6 figs. 

1948. Annehda, Polychaeta errantia. In Canadian Pacific Fauna. Fish. 
Res. Board Canada, No. 9b (1), pp. 1-100, 160 figs. 

1950. Notes on Polychaeta from the coast of western Canada. IV. Poly- 
chaeta sedentaria. Ann. Mag. Nat. Hist., ser. 12, vol. 3, No. 25, 
pp. 50-69, 8 figs. 

1952. Annelida, Polychaeta sedentaria. In Canadian Pacific Fauna. 
Fish. Res. Board Canada, No. 9b (2), pp. 1-139, 292 figs. 

BORG, FOLKE 

1917. t)ber die Spirorbisarten Schwedens nebst einem Versuch zu einer 
neuen Einteilung der Gattung Spirorbis. Zool. Bidr. Uppsala, 
vol. 5, pp. 14-48, 16 figs. 
Chamberlin, Ralph V. 

1919. Pacific coast Polychaeta collected by Alexander Agassiz. Bull. Mus. 

Comp. Zool., vol. 63, No. 6, pp. 251-276, pis. 1-3. 

1920. The polychaetes collected by the Canadian Arctic Expedition, 

1913-18. Rep. Canadian Arctic Exped. 1913-18, pt. B, vol. 9, 
pp. 1-41, 6 pis. 

CLAPARfeoE, finOUARD 

1868. Les ann^lides ch^topodes du Gulfe de Naples. M6m. Soc. Phys. 
Genfeve, vol. 19, pt. 2, pp. 313-584, 16 pis. 
261112—54 10 



348 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Dalyell, John Graham 

1853. The powers of the Creator displayed in the Creation. Vol. 2, pp. 1- 
359, 46 pis. 

DiTLEVSBN, HjALMAR 

1909. Annulata Polychaeta. In Report of the second Norwegian Arctic 

expedition in the Fram, 1898-1902, vol. 3, No. 15, pp. 1-23, 3 pis. 
1911. Annelids from the Danmark Expedition. In Danmark-Ekspedi- 

tionen til Gr0nlands Nord0stkyst 1906-1908, vol. 5, No. 9, pp. 411- 

432, pis. 27-31. 
1917. Annelids. I. In The Danish 7n^oZ/-Expedition, vol. 4, pt. 4, 

pp. 1-71, 24 figs., 6 pis. 
1929. Polychaeta. In Zoology of the Faroes. No. 16, pp. 1-83. 
1937. Polychaeta. The Godthaab Expedition, 1928, vol. 80, No. 4, 

pp. 1-64, 6 figs. 
Ehlers, Ernst 

1868. Die Borstenwiirmer (Annelida Chaetopoda) nach systematischen 

und anatomischen untersuchungen dargestellt, vol. 2, pp. 269-748, 

pis. 12-24. 

1913. Die Polychaeten-Sammlungen. In Deutschen Siidpolar-Expedition, 

1901-1903, Zool., vol. 5, pt. 4, pp. 399-602, pis. 26-46. 
Eliason, Anders 

1920. Biologisch-Faunistische Untersuchungen aus dem Oresund. V. 
Polychaeta. Acta Univ. Lundensis, new ser., vol. 16, pt. 6, 
pp. 1-103, 18 figs. 
Fabricius, Otto 

1780. Fauna groenlandica. . . . Pp. 1-452, 12 figs. 

1799. Betragtninger over Nereide-slaegten. Skrivter af Naturh. Selsk., 
vol. 5, pp. 154-190, pi. 4. 
Fauvel, Pierre 

1911. Ann^lides polych&tes. In Campagne Arctique de 1907, Due d' Or- 
leans, pt. 9, pp. 1-44, 1 pi. 

1914. Ann(51ides polychStes non p^lagiques provenant des campagnes de 

VHirondelle et de la Princesse- Alice (1885-1910), fasc. 46, pp. 1- 

432, 31 pis. 
1923. Polych^tes errantes. Faune de France, vol. 5, pp. 1-488, 181 figs. 
1927. Polychfetes s6dentaires. Faune de France, vol. 16, pp. 1-494, 152 

figs. 

1932. Annelida polychaeta of the Indian Museum, Calcutta. Mem. 

Indian Mus., vol. 12, No. 1, pp. 1-262, 40 figs., 9 pis. 

1933. Ann^lides polych5tes du Golfe du Pei Tcheu Ly. Publ. Mus. Hoang 

ho Pai ho de Tien Tsin, No. 15, pp. 1-67, 6 figs. 
1934a. Ann^lides polych^tes de Rovigno d'Istria. Thalassia, vol. 1, No. 7, 

pp. 1-78, 4 figs. 
1934b. Sur quelques syllidiens du Japon. Annot. Zool. Japon., vol. 14, 

No. 3, pp. 301-315, 2 figs. 
1936. Ann^lides polychfites du Japon. Mem. Coll. Sci. Kyoto Univ., 

ser. B, vol. 12, No. 1, pp. 41-92, 1 fig. 
1947. Ann^Iides polychfetes de Nouvelle-Cal^donie et des lies Garabier. 

In Faune de I'Empire Francais, vol. 8, pp. 1-107, 90 figs. 
Friedrich, Hermann 

1939. Polychaeten-Studien, IV. Zur Polychaeten fauna der Barents-See. 

Kieler Meeresf., vol. 3, pt. 1, pp. 122-132, 6 figs. 



MARINE POLYCHAETE WORMS — PETTIBONE 349 

GORBUNOV, G. P. 

1946. Bottom life of the Novosiberian shoalwaters and the central part 
of the Arctic ocean. Trudy DreifufBshchei ekspeditsii Glavsev- 
morputi na ledokolnom parokhode G. Sedov 1937-1940, vol. 3, 
pp. 30-138, 1 pi. (Leningrad Vsesoiuznyl arkticheskil institut). 
Grube, Adolph Eduard 

1840. Actinien, Echinodermen und Wiirmen des Adriatischen und Mit- 
telmeers. ... Pp. 61-88, 1 pi. 

1855. Beschreibungen neuer oder wenig bekannter Anneliden. Arch. 
Naturg., vol. 21, pt. 1, pp. 81-136, pis. 3-5. 

1860. Beschreibung neuer oder wenig bekanter Anneliden. Arch. Naturg., 
vol. 26, pt. 1, pp. 71-118, pis. 3-5. 

GXJSTAFSON, GUNNAR 

1936. Polychaeta and Sipunculoidea from the Siberian Arctic ocean. In 

The Norwegian North Polar expedition with the Afaud, 1918-1925, 

Scientific Results, vol. 5, No. 17, pp. 1-12. 
Hansen, G. Armauer 

1882. Anneliden fra den Norske Nordhavs-Expedition, 1876-1878. Den 

Norske Nordhavs-Exped., pt. 7, pp. 1-53, 7 pis. 
Hartman, Olga 

1936a. A review of the Phyllodocidae (Annelida Polychaeta) of the coast of 

California, with descriptions of nine new species. Univ. Calif. 

Publ. Zool., vol. 41, No. 10, pp. 117-132, 51 figs. 
1936b. New species of polychaetous annelids of the family Nereidae from 

California. Proc. U. S. Nat. Mus., vol. 83, No. 2994, pp. 467-480, 

figs. 46-53. 
1936c. New species of Spionidae (Annelida Polychaeta) from the coast of 

California. Univ. Calif. Publ. Zool., vol. 41, No. 6, pp. 45-52, 

22 figs. 
1938a. The types of the polychaete worms of the familes Polynoidae and 

Polyodontidae in the United States Museum and the description 

of a new genus. Proc. U. S. Nat. Mus., vol. 86, pp. 107-134, 

41 figs. 
1938b. Review of the annelid worms of the family Nephtyidae from the 

northeast Pacific, with descriptions of five new species. Proc. 

U. S. Nat. Mus., vol. 85, No. 3034, pp. 143-158, figs. 62-67. 
1938c. Annotated list of the types of polychaetous annelids in the Museum 

of Comparative Zoology. Bull. Mus. Comp. Zool., vol. 85, No. 1, 

pp. 1-31, 3 pis. 

1939. Polychaetous annehds. Pt. I. Aphroditidae to Pisionidae. Allan 

Hancock Pacific Exped., vol. 7, No. 1, pp. 1-156, 28 pis. 

1940. Polychaetous annelids. Pt. II. Chrysopetalidae to Goniadidae. 

Allan Hancock Pacific Exped., vol. 7, No. 3, pp. 173-286, pis. 
31-44. 

1941a. Some contributions to the biology and life history of Spionidae 
from California. Allan Hancock Pacific Exped., vol. 7, No. 4, 
pp. 289-322, pis. 45-48. 

1941b. Polychaetous annelids. Pt. IV. Pectinariidae. Allan Hancock Pa- 
cific Exped., vol. 7, No. 5, pp. 325-344, pis. 49-52. 

1942a. A review of the types of polychaetous annelids at the Peabody 
Museum of Natural History, Yale University. Bull. Bingham 
Oceanogr. Coll., vol. 8, pt. 1, pp. 1-98, 161 figs. 



350 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

1942b. The identity of some marine annelid worms in the United States 

National Museum. Proc. U. S. Nat. Mus., vol. 92, No. 3142, pp. 

101-140, figs. 8-15. 
1942c. Report on the scientific results of the Atlantis expeditions to the 

West Indies under the joint auspices of the University of Havana 

and Harvard University. Mem. Soc. Cubana Hist. Nat., vol. 16, 

No. 2, pp. 89-104, pis. 8-9. 
1944a. New England Annelida. Pt. 2, including the unpublished plates by 

Verrill with reconstructed captions. Bull. Amer. Mus. Nat. Hist., 

vol. 82, No. 7, pp. 327-344, pis. 13-35 [45-60]. 
1944b. Polychaetous annelids from California including the descriptions of 

two new genera and nine new species. Allan Hancock Pacific 

Exped., vol. 10, No. 2, pp. 239-307, pis. 19-26. 
1944c. Polychaetous annelids. Pt. VI. Paraonidae, Magelonidae, Longo- 

somidae, Ctenodrilidae, and Sabellariidae. Allan Hancock Pacific 

Exped., vol. 10, No. 3, pp. 311-388, pis. 27-42. 
1944d. Polychaetous annelids. Allan Hancock Atlantic Exped., Rep. No. 

3, pp. 1-32, 2 pis. 
1945. The marine annelids of North Carolina. Bull. Duke Univ. Marine 

Sta., No. 2, pp. 1-51, 10 pis. 

1947. Polychaetous annelids. Pt. VII. Capitellidae. Allan Hancock Pa- 

cific Exped., vol. 10, No. 4, pp. 391-480, pis. 43-58. 

1948. The polychaetous annelids of Alaska. Pacific Sci., Univ. Hawaii, 

vol. 2, No. 1, pp. 3-58, 12 figs. 

1950. Goniadidae, Glyceridae and Nephtyidae, Allan Hancock Pacific 

Exped., vol. 15, No. 1, pp. 1-180, 19 pis., 3 figs. 

1951. The littoral marine annelids of the Gulf of Mexico. Publ. Inst. 

Marine Sci., vol. 2, No. 1, pp. 7-124, 27 pis. 

1952. The marine annelids of the United States Navy Antarctic Expedi- 

tion, 1947-1948. Journ. Washington Acad. Sci., vol. 42, No. 7, 
pp. 231-237, 12 figs. 
Hartman, Olga, and Reish, Donald J. 

1950. The marine annelids of Oregon. Oregon State Monogr. Studies in 
Zool., No. 6, pp. 1-64, 5 pis. 
Hessle, Christian 

1917. Zur Kenntnis der Terebellomorphen Polychaeten. Zool. Bidr 
Uppsala, vol. 5, pp. 39-258, 66 figs,, pis. 1-5, 
Johansson, Karl Erik 

1926. Berkmurkungen uber die Kinberg'schen Arten der Familien Hermelli- 

dae und Sabellidae. Ark. Zool. Stockholm, vol. 18A, No. 7, pp. 
1-28, 9 figs. 

1927. Beitrage zur Kenntnis der Polychaeten Familien Hermellidae, 

Sabellidae und Serpulidae. Zool. Bidr, Uppsala, vol. 11, pp. 1-184, 

15 figs., 5 pis. 
Johnson, Herbert Parlin 

1897. A preliminary account of the marine annelids of the Pacific coast with 

descriptions of new species. . . . Proc. Calif. Acad. Sci. Zool,, 

ser, 3, vol. 1, No. 5, pp. 153-198, pis. 5-10. 
1901. The Polychaeta of the Puget Sound region. Proc. Boston Soc. Nat. 

Hist., vol. 29, No. 18, pp. 381-437, 19 pis. 

KiNBERG, J. G. H. 

1867. Annulata nova. Ofv. Vet. Akad. Forh. Stockholm, vol. 23, No. 9, 
pp. 337-357. 



MARINE POLYCHAETE WORMS — PETTIBONE 351 

Kroyer, Henrik 

1856. Meddelelser om ormeslaegten Sabella Linn., isaer dans nordiske Arter. 
Overs. Danske Videnskab. Selsk. Forh. (1856), pp. 1-36. 
Langerhans, Paxil 

1884. Die Wurmfauna von Madeira. IV. Zeits. wiss. Zool., vol. 40, pp. 

247-285, pis. 15-17. 
Leuckart, Rudolph 

1849. Zur Kenntniss der Fauna von Island. Erster Beitrag (Wurmer). 
Arch. Naturg., vol. 15.1, pp. 149-208, pi. 3. 
Levinsen, G. M. R. 

1882. Systematisk-geografisk Oversigt over de nordiske Annulata, Gephyrea, 
Chaetognathi og Balanoglossi. Vidensk. Medd. Naturh. Foren. 
Kj0benhavn, pt. 1, pp. 160-251, pi. 7. 
LiNNE, Carl 

1758. Systema Naturae. Ed. 10. 
1767. Systema Naturae. Ed. 12. 
McIntosh, William C. 

1874. On the AnneUda of the Gulf of St. Lawrence. Ann. Mag. Nat. Hist., 
ser. 4, vol. 13, pp. 261-270, pis. 9-10. 

1885. Annelida Polychaeta. In Report on the scientific results of the voyage 

of H.M.S. Challenger . . . 1873-76 . . . Zoology, vol. 12, pt. 34, 

pp. 1-554, 84 pis. 
1900. A monograph of the British annelids. II. Polychaeta. Amphino- 

midae to Sigalionidae. Vol. 1, pp. 215-442, pis. 24-42. 
1908. A monograph of the British annelids. I-II. Polychaeta. Neph- 

thydidae to Ariciidae. Vol. 2, pp. 1-524, pis. 43-87. 
Malm, August W, 

1874. Annulater i hafvet utmed Sverges vestkust och omkring Goteborg. 

Goteborgs Vetensk. Samh. Handl., vol. 14, pp. 71-105, 1 pi. 
Malmgeen, Anders J. 

1865. Nordiska Hafs-Annulater. Forh. Of v. Kongl. Vet, Akad. Stockholm, 

Nos. 1, 2, 5, pp. 51-110, 181-192, 355-410, pis. 8-15, 18-29. 
1867. Annulata Polychaeta Spetsbergiae, Groenlandiae, Islandiae et Scan- 

dinaviae hactenus cognita. Pp. 1-127, 14 pis. 
Marenzeller, Emil von 

1876. Zur Kenntniss der adriatischen Anneliden. Sitzb. Akad. Wiss. Wien, 

vol. 72, pp. 129-171, 4 pis. 
1879. Siidjapanische AnneHden. Denkschr. Akad. Wiss. Wien, vol. 41, pp. 

1-46, 6 pis. 
1890. Annulaten des Beringmeeres. Ann. Naturh. Hofmus. Wien, vol. 5, 

pp. 1-8, 1 pi. 
1892. Zoologische Ergebnisse der im Jahre 1889 auf Kosten der Bremer 

Geographischen Gesellschaft von Dr. Willy Kiikenthal und Dr. 

Alfred Walter ausgefiihrten Expedition nach Ostspitzbergen. 

Polychaten. Zool. Jahrb., vol. 6, pp. 397-434, pi. 19. m 

1904. Zoologische Ergebnisse. XIII. Polychaten des Grundes, Gesam- 

melt 1893, 1894. Denkschr. Akad. Wiss. Wien, vol. 74," pp. 

295-321. 
Mesnil, Ff;Lix j«:i 

1897. fitudes de morphologie externe chez les ann^lides. II. Remarques 

complementaires sur les spionidiens. Bull. Sci. France et Belgique, 

vol. 30, pp. 83-100, pi. 3. 



352 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Monro, C. C. A. 

1928. On the Polychaeta collected by Dr. Th. Mortensen off the coast of 

Panama. Vidensk. Medd. Naturh. Foren. Kj0benhavn, vol. 85, 

pp. 75-103, 19 figs. 
1930. Polychaete worms. Discovery Reports, vol. 2, pp. 1-222, 91 figs. 
1933. The Polychaeta Errantia collected by Dr. C. Crossland at Col6n in 

the Panama region and Galdpagos Islands during the expedition 

of the S.Y. St. George. Proc. Zool. Soc. London (1933), pt. 1, pp. 

1-96, 36 figs. 

1936. Polychaete worms. II. Discovery Reports, vol. 12, pp. 59-198, 34 

figs. 

1937. Polychaeta. Scientific reports of the John Murray Expedition, 

1933-34, vol. 4, No. 8, pp. 243-321, 28 figs. 
1939a. Polychaeta of the Rosaura Expedition. Nov. Zool. London, vol. 

41, pp. 345-354, 4 figs. 
1939b. On some tropical Polychaeta in the British Museum, mostly collected 

by Dr. C. Crossland at Zanzibar, Tahiti and the Marquesas. II. 

Families Syllidae and Hesionidae. Nov. Zool. London, vol. 41, 

pp. 383-393, 4 figs. 
Montagu, George 

1808. Description of several marine animals found on the south coast of 

Devonshire. Trans. Linn. Soc. London, Zool., vol. 9, pp. 81-114, 

pis. 2-8. 
1818. Description of five British species of the genus Terebella of Linn4. 

Trans. Linn. Soc. London, Zool., vol. 12, pp. 340-344, pis. 11-13. 
Moore, J. Percy 

1902. Descriptions of some new Polynoidae with a list of other Polychaeta 

from North Greenland waters (and Alaska). Proc. Acad. Nat. 
Sci. Philadelphia, vol. 54, pp. 258-278, pis. 13-14. 

1903. Polychaeta from the coastal slope of Japan, and from Kamchatka 

and Bering Sea. Proc. Acad. Nat. Sci. Philadelphia, vol. 55, pp. 

401-490, pis. 23-27. 
1905a. New species of Polychaeta from the North Pacific, chiefly from 

Alaskan waters. Proc. Acad. Nat. Sci. Philadelphia, vol. 57, pp. 

525-554, pis. 34-37. 
1905b. New species of Ampharetidae and Terebellidae from the North 

Pacific. Proc. Acad. Nat. Sci. Philadelphia, vol. 57, pp. 846-860, 

pi. 44. 
1905c. Five new species of Pseudopotamilla from the Pacific coast of North 

America. Proc. Acad. Nat. Sci. Philadelphia, vol. 57, pp. 555- 

569, pi. 37. 
1906a. Descriptions of two new Polychaeta from Alaska. Proc. Acad. Nat. 

Sci. Philadelphia, vol. 58, pp. 352-355, 2 figs. 
1906b. Additional new species of Polychaeta from the North Pacific. Proc. 

Acad. Nat. Sci. Philadelphia, vol. 58, pp. 217-260, pis. 10-12. 
1908. Some polychaetous annelids of the northern Pacific coast of North 

America. Proc. Acad. Nat. Sci. Philadelphia, vol. 60, pp. 321- 

364, 4 figs. 
1909a. The polychaetous annelids dredged by the U.S.S. Albatross off the 

coast of southern California in 1904. I. Syllidae, Sphaerodoridae, 

Hesionidae and Phyllodocidae. Proc. Acad. Nat. Sci. Phila- 
delphia, vol. 61, pp. 321-351, pis. 15-16. 



MARINE POLYCHAETE WORMS — PETTIBONE 353 

1909b. The polychaetous annelids dredged in 1908 by Mr. Owen Bryant 
oflf the coasts of Labrador, Newfoundland, and Nova Scotia. 
Proc. U. S. Nat. Mus., vol. 37, No. 1703, pp. 133-146. 

1910. The polychaetous annelids dredged by the U.S.S. Albatross oflf the 

coast of southern Cahfornia in 1904. II. Polynoidae, Aphro- 
ditidae and Segaleonidae. Proc. Acad. Nat. Sci. Philadelphia, 
vol. 62, pp. 328-402, pis. 28-33. 

1911. The polychaetous annelids dredged by the U.S.S. Albatross off the 

coast of southern California in 1904. Ill, Euphrosynidae to 

Goniadidae. Proc. Acad. Nat. Sci. Philadelphia, vol. 63, pp. 

234-318, pis. 15-21. 
1923. The polychaetous annehds dredged by the U.S.S. Albatross off the 

coast of Southern California in 1904. IV. Spionidae to Sabel- 

lariidae. Proc. Acad. Nat. Sci. Philadelphia, vol. 75 (Append.), 

pp. 179-259. 
Moore, J. P., and Bush, K. J. 

1904. Sabellidae and Serpulidae from Japan, with descriptions of new 

species of Spirorbis. Proc. Acad. Nat. Sci. Philadelphia, vol. 

56, pp. 157-179, pis. 11, 12. 
MtJLLER, Otto Frederich 

1776. Zoologiae Danicae prodromus, seu animalium Daniae et Norvegiae 

. . ., pp. 1-274. 
1788. Zoologica Danica, seu animalium Daniae et Norvegiae . . ., vol. 

1, ed. 3, pp. 1-52; vol. 3, ed. 3, pp. 1-71, 1789. 
1806. Atlas to above volumes. 

Murdoch, John 

1884. Description of seven new species of Crustacea and one worm from 

Arctic Alaska. Proc. U. S. Nat. Mus., vol. 7, pp. 518-522, 

1885, Natural history of Alaska, Vermes. In Report of the Inter- 

national Polar Expedition to Point Barrow, Alaska, pt. 4, pp. 
152-156. 
NiLssoN, David 

1928. Neue und alte Amphicteniden. Goteborgs Vetensk. Samh, Handl,, 
vol, 33, No. 4, pp. 1-96, 30 figs. 
Oersted, Anders S. 

1843. Groenlands Annulata Dorsibranchiata. K. Danske Videns. Selsk. 

Naturv, Math., vol. 10, pp. 155-216, 8 pis, 
1845. "Qber die Entwickelung der Jungen bei einer Annelide und iiber die 
atisseren Unterschiede zwischen beiden Geschlechtern. Arch. 
Naturg., vol. 11.1, pp. 20-23, pi. 2. 
Okuda, Shiro 

1937a. Some ariciid worms from Japan. Annot. Zool. Japon., vol. 16, No. 

2, pp. 99-105, 6 figs, 

1937b. Annelida Polychaeta in Onagawa Bay and its vicinity. I. Polychaeta 
Sedentaria. Sci. Rep. T6hoku Univ., ser. 4, vol, 12, No. 1, pp. 
45-69, 12 figs., pi. 2. 

1938a, The Sabellariidae of Japan. Journ. Fac. Sci. Hokkaido Univ. Zool,, 
ser. 6, vol. 6, No. 3, pp. 235-253, 11 figs. 

1938b. Polychaetous annelids from the vicinity of the Mitsui Institute of 
Marine Biology. Japan. Journ. Zool., vol. 8, No. 1, pp. 75-105, 
15 figs, 

1939, Annelida Polychaeta in Onagawa Bay and its vicinity, II, Poly- 
chaeta Errantia. Sci. Rep. Tohoku Univ., ser. 4, vol. 14, No. 2-3, 
pp. 219-244, 14 figs. 



354 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m 

Pagenstecher, H. a. 

1862. Untersuchungen uber niedere Seethiere aus Cette. I. Exogone gem- 
mifera und einige verwandte Syllidien. Zeits. Wiss. ZooL, vol. 12, 
pp. 265-283, pis. 25-26. 
Pallas, Peter Simon 

1766. Miscellanea zoologica quibus novae imprimis atque obscurae ani- 
malium species describuntur et observationibus inconibusque illus- 
trantur. Pp. 1-244, 14 pis. 
Pettibone, Marian H. 

1949. Polychaetous annelids of the Polynoidae from the northeastern 
Pacific, with a description of a new species. Amer. Mus. Nov., 
No. 1414, pp. 1-5, 9 figs. 
1951. A new species of polychaete worm of the family Polynoidae from 
Point Barrow, Alaska. Journ. Washington Acad. Sci., vol. 41, 
No. 1, pp. 44-45, 1 fig. 
1953. Some scale-bearing polychaetes of Puget Sound and adjacent waters 
Pp. 1-89, 40 pis. 
Ranzani, Camillo 

1817. Neue Wtirmer. Isis von Oken, vol. 1, pp. 1449-1480, pi. 11. 
Rathke, Heinrich 

1843. Beitrage zur Fauna Norwegens. Nova Acta Acad. Leop. Carol., 
vol. 20, No. 1, pp. 1-264, 12 pis. 
RiojA, Enrique 

1941. Estudios annelidol6gicos. III. Datos para el conocimiento de las 
fauna de poliquetos de las costas del Pacffico de Mexico. Anal. 
Inst. Biol. Mexico, vol. 12, No. 2, pp. 669-746, 9 pis. 
1943. Estudios annelidol6gicos. VIII. Aportaciones al conocimiento de los 
exogoninos (anel. poliquetos) de las costas Mexicanas del Pa- 
cffico. Anal. Inst. Biol. M6xico, vol. 14, No. 1, pp. 207-227, 47 
figs. 
Sars, Michael 

1835. Beskrivelser og iagttagelser over nogle moekelige eller nye i havet ved 

den Bergenske kyst levende dyr. . . . Pp. 1-81, 15 pis. 
1851. Beretning om en i Sommeren 1849 foretagen zoologisk Reise i Lofoten 
og Finmarken. 5. Annelider. Nyt. Mag. Naturv., vol. 6, pp. 
196-211. 
1860. Om de ved Norges Kyster forekommende Arter af Annelides laegten 
Polynoe. Vidensk. Selsk. Kristiania, pp. 54-62. 
Smith, S. I., and Harger, O. (and Verrill, A. E.) 

1874. Report on the dredgings in the region of St. George's Banks in 1872. 
Trans. Connecticut Acad. Arts Sci., vol. 3, pp. 1-57, 8 pis. 
Southern, Rowland 

1914. Archiannelida and Polychaeta. In Clare Island survey* Pt. 47. 
Proc. Roy. Irish Acad. Dublin, vol. 31, pp. 1-160, 15 pis. 
Stimpson, William 

1854. Synopsis of the marine Invertebrata of Grand Manan. Smithsonian 
Contr. Knowl., vol. 6, pp. 1-66, 3 pis. 
St0p-Bowitz, C. 

1941. Les glyc^riens de Norvfege. Nyt. Mag. Naturv., vol. 82, pp. 181-250 

4 pis., 14 figs. 
1946. Les scalibregmiens de Norvfege. Nyt. Mag. Naturv., vol. 85, pp. 
63-87, 2 pis., 6 figs. 



MARINE POLYCHAETE WORMS — PETTIBONB 



355 



1948a. Les flabelligeriens Norv^giens. Bergens Mus. Aarb. 1946/47, 

Naturv., No. 2, pp. 1-59, 13 figs., 12 pis. 
1948b. Sur les polychfetes arctiques des families des glyc^riens, des oph61iens, 

des scalibregmiens et des flabelligeriens. Troms0 Mus. Aarsh., 

vol. 66, No. 2, pp. 1-58, 18 figs. 
1948c. Polychaeta from the Michael Sars North Atlantic deep-sea expedition 

1910. In Rep. Sci. Results Michael Sars North Atlantic Deep-sea 

Exped. 1910, vol. 5, No. 8, pp. 1-91, 51 figs., 5 tables. 
Sumner, F. B., Osburn, R. C, and Cole, L. J. 

1913. Annulata. In A biological survey of the waters of the Woods Hole 

and vicinity. Bull. U. S. Bur. Fish., vol. 31, pt. 2, pp. 615-636. 
Th6el, Johan H. 

1879. Les annelides polychfetes des mers de la Nouvelle-Zemble. Svenska 

Vetensk. Akad. Handl., vol. 16, pt. 3, pp. 1-75, 4 pis. 
Thorson, Gunnar 

1946. Reproduction and larval development of Danish marine bottom 

invertebrates. Medd. Komm. Havunders0g. Kj0benhavn, vol. 4, 

No. 1, pp. 1-523, 198 figs. 
Treadwelx,, a. L. 

1922. Polychaetous annelids collected at Friday Harbor, State of Wash- 
ington, in February and March, 1920. Publ. Carnegie Inst., 

No. 312, pp. 171-181, 37 figs. 
1925. A list of the annelids collected by Captain R. A, Bartlett in Alaska, 

1924, with a description of a new species. Proc. U. S. Nat. Mus., 

vol. 67, art. 29, No. 2601, pp. 1-3, 4 figs. 
1937. Polychaetous annelids collected by Captain Robert A. Bartlett in 

Greenland, Fox Basin, and Labrador. Journ. Washington Acad. 

Sci., vol. 27, No. 1, pp. 23-36, 16 figs. 
Verrill, Addison Emory 

1873. Report upon the invertebrate animals of Vineyard Sound and the 

adjacent waters, with an account of the ph}'sical characters of the 

region. Rep. U. S. Fish Comm. for 1871-72, pp. 295-778. 
1874a. Brief contributions to zoology from the Museum of Yale College. 

Results of recent dredging expeditions on the coast of New England. 

Amer. Jour. Sci. Arts, vol. 7, No. 4-8, pp. 38-46, 131-138, 405- 

414, 498-505, pis. 4-8. 
1874b. Explorations of Casco Bay by the U. S. Fish Commission in 1873. 

Proc. Amer. Assoc. Adv. Sci., vol. 22, pp. 340-395, 6 pis. 
1879. Notice of recent additions to the marine invertebrata of the north- 
eastern coast of America, with descriptions of new genera and 

species and critical remarks on others. Proc. U. S. Nat. Mus., 

vol. 2, pp. 165-206. 

1881. New England Annelida. Pt. I. Historical sketch, with annotated 

lists of the species hitherto recorded. Trans. Connecticut Acad. 
Arts Sci., vol. 4, pt. 2, pp. 285-324, pis. 3-12. 

1882. Notice of the remarkable marine fauna occupying the outer banks 

off the southern coast of New England, and of some additions to 
the fauna of Vineyard Sound. Amer. Journ. Sci., ser. 3, vol. 24, 
pp. 360-371 (plates referred to published in Hartman, 1944a). 
Webster, H. E. 

1879a. Annelida Chaetopoda of the Virginia coast. Trans. Albany Inst., 
vol. 9, pp. 201-272, 11 pis. 



356 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

1879b. The Annelida Chaetopoda of New Jersey. Rep, New York State 
Mus., vol. 32, pp. 101-128. 

1886. The Annelida Chaetopoda of New Jersey. Rep. New York State 

Mus., vol. 39, pp. 128-159, pis. 4-10. 
Webster, H. E., and Benedict, J. E. 

1884. The Annelida Chaetopoda from Provincetown and Wellfleet, Massa- 
chusetts. Rep. U. S. Fish Comm. for 1881, pp. 699-747, 8 pis. 

1887. The Annelida Chaetopoda from Eastport, Maine. Rep. U. S. Fish 

Comm. for 1885, pp. 707-755, 8 pis. 
Wesbnberg-Lund, Elise 

1934. Gephyreans and annelids. In The Scoresby Sound Committee's 

second East Greenland expedition in 1932 to King Christian IX's 

Land. Medd. Gr0nland, vol. 104, No. 14, pp. 1-38. 
1939a. Pelagic polychaetes of the families Aphroditidae, Phyllodocidae, 

Typhloscolecidae and Alciopidae. II. Biology. In Report on the 

Danish oceanographical expeditions 1908-10 to the Mediterranean 

and adjacent seas, pp. 1-46, 29 figs. 
1939b. Polychfetes et g^phyriens de Tunisie. Bull. Stat. Oc^anogr. Salambo, 

No. 39, pp. 1-20. 

1947. Syllidae (Polychaeta) from Greenland waters. Medd. Gr0nland, 

vol. 134, No. 6, pp. 1-38, 15 figs. 

1948. Maldanidae (Polychaeta) from west Greenland waters. Medd. 

Gr0nland, vol. 134, No. 9, pp. 1-58, 29 figs. 

1949. Polychaetes of the Iranian Gulf. Danish Sci. Invest. Iran, pt. 4, 

pp. 247-400, 47 figs. 
1950a. Polychaeta. In The Danish Zn^/oZf-Expedition, vol. 4, No. 14, pp. 1-92, 

10 pis., 67 charts, 2 figs. 
1950b. The Polychaeta of west Greenland. . . . Medd. Gr0nland, vol. 151, 

No. 2, pp. 1-171, 37 charts, 4 tables. 
1951. Polychaeta. In The Zoology of Iceland, vol. 2, pt. 19, pp. 1-182, 

12 figs., 62 charts. 

WlE^N, AXBL 

1883. Chaetopoder fr&n Sibiriska Ishafvet och Berings Haf insamlade 

under Feg^a-Expeditionen 1878-1879. In Fe^a-Exped. veten- 

skapliga lakttagelser. . . . af A. E. Nordenskiold, vol. 2, pp. 

383-428, pis. 27-32. 
1901. Uber die wahrend der schwedischen arktischen Expedition von 1898 

und 1900 eingesammelten Anneliden. Zool. Anz., vol. 24, p. 253. 
Zachs, I. 

1933. Polychaeten vom Nordjapanischen Meer (Bucht Peter der Grosse). 

Issled. Morei USSR, Gosud. Gidrol. Inst., Leningrad, fasc. 19, 

pp. 125-137. 
Zatsepin, V. 

1948. Klass Polychaeta. In Opredelitel fauny i flory severnykh morel 

USSR, N. S. Gaevskaia-Sokolova, pp. 94-167, figs. 18-22, pis. 28-39. 



U. 3. GOVERNMENT PRINTING OFFICEi I9B4 



PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 




issued 



SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 



Vol. 103 Washington : 1954 No. 3325 

THE RELATIONSHIPS OF OLD AND NEW WORLD 
MELANIANS 



By J. P. E. Morrison 



Recent anatomical observations on the reproductive systems of 
certain so-called "melanian" fresh-water snails and their marine rela- 
tives have clarified to a remarkable degree the supergeneric relation- 
ships of these fresh-water forms. 

The family of Melanians, in the broad sense, is a biological ab- 
surdity. We have the anomaly of one fresh-water "family" of snails 
derived from or at least structurally identical in peculiar animal 
characters to and ancestrally related to three separate and distinct 
marine famiHes. On the other hand, the biological picture has been 
previously misunderstood largely because of the concurrent and 
convergent evolution of the three fresh-water groups, Pleuroceridae, 
Melanopsidae, and Thiaridae, from ancestors common to the marine 
families Cerithiidae, Modulidae, and Planaxidae, respectively. 

The family Melanopsidae is definitely known living only in Europe. 
At present, the exact placement of the genus Zemelanopsis Uving in 
fresh waters of New Zealand is uncertain, since its reproductive 
characters are as yet unknown. In spite of obvious differences in 
shape, the shells of the marine genus Modulus possess at least a well- 
indicated columellar notch of the aperture, to corroborate the biologi- 
cal relationship indicated by the almost identical female egg-laying 
structure in the right side of the foot of Modulus and Melanopsis. 



273553—54 1 357 



358 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

The family Pleuroceridae, fresh-water representative of the ancestral 
cerithiid stock, is now known to include species living in Africa, 
Asia, and the Americas. 

The family Thiaridae, likewise, is known from Africa, Asia, and 
America. With identical reproductive characters, the Thiaridae are 
only too obviously the fresh-water relatives of the Planaxidae. It is 
only when the Melanopsidae, Pleuroceridae, and Thiaridae are 
separated that their zoogeographic story can be read without confu- 
sion. Perfect separation of these three fresh-water families can only 
be accomplished by reference to characters of the animals, particularly 
the reproductive characters. Radular, opercular, or shell characters 
that will completely separate these families are nonexistent because of 
their concurrent and convergent evolution. 

The following key to the major groups of the Melanian complex is 
given here as the most logical one possible. In the author's opinion, 
it represents the true biological relationship of these groups. 



c* Reproduction dioecious; males present in species. 

6.1 Female with large grooved "ovipositor" in pit on right side of foot; few eggs 

of proportionately large size. 

c' Marine shells Modulidae 

C.2 Fresh-water shells Melanopsidae 

6.2 Female with egg-laying sinus and rudimentary papilla, or no special struc- 

ture on right side of foot; numerous eggs of proportionately smaller size. 

d.i Marine shells Cerithiidae 

d.^ Fresh-water shells Pleuroceridae 

c.i Males with no intromittent structures. 
/.I Females oviparous, with egg-laying sinus in right side of foot. 

Pleurocerinae 
/.2 Females ovo viviparous, without egg-laying sinus; brood pouch 

uterine, the enlarged end of the oviduct Lavigeriinae 

e.^ Males •with eversible "penis" in mantle edge near end of vas deferens; 

females ovo viviparous; brood pouch uterine Tiphobiinae 

a.2 Reproduction parthenogenetic; no males present in species; brood pouch not 
uterine, but adventitious (subhaemocoelic) in the neck region, with opening 
on right side of neck. 

g.^ Marine shells Planaxidae 

0.2 Fresh- water shells Thiaridae 



Family Melanopsidae (Europe) 

Two genera from the fresh waters of Europe have been studied and 
proven to belong to the family Melanopsidae. They are more special- 
ized than the Asiatic and American Pleuroceridae, possessing a much 
larger "ovipositor" and laying a much smaller number of proportion- 
ately much larger eggs. 



OLD AND NEW WORLD MELANIANS — MORRISON 359 

According to studies by Ankel (1928), Fagotia esperi (Ferussac) 
has the same reproductive anatomy of the female as does Melanopsis. 
The few large eggs (1 mm. in size) are laid singly in an irregular capsule. 

Melanopsis dufourii (Ferussac) from Elche, Alicante Province, 
Spain, has been examined by me. Unlike the animals of the North 
American Pleuroceridae, in this species at least, the smooth mantle 
edge is continuous beneath the back of the foot, forming a circle around 
the aperture. In the female the end of the oviduct is quite far behind 
the mantle edge and simple, as is typical for the group. The egg- 
laying groove is not long, and does not even reach the mantle edge. 
The pit is enormously developed and has a large, whitish, tongue- 
shaped, basally-attached "ovipositor," with a gi'oove on its posterior 
dorsal face, toward the rest of the pit. According to Ankel (1928), 
a few proportionately large eggs are laid by Melanopsis. 

Family Pleuroceridae (in the Americas) 
Subfamily pleurocerinae 

All known American members of Pleuroceridae are oviparous, 
including those from North, Central, and South America and the 
West Indies. In other words, every American genus and species for 
which the characters of reproduction are known is egg-laying, with an 
egg-laying sinus or pit in the right side of the foot of the females. The 
presence of this pit is the single morphological character that proves 
them to belong to the typical subfamily. There is every indication 
that all the American genera and species of Pleuroceridae belong here, 
although some of the North American genera are still unrecorded as to 
critical animal characters. 

Subfamily pleurocerinae (in North America) 
Genus Oxytrema Rafinesque, 1819 

Oxytrema Rafinesque is the earliest and correct name for one of the 
most widespread "Melanian" genera in the world. This genus in- 
cludes numerous North American species whose ranges extend from 
the Atlantic to the Pacific coasts and from southern Canada to Florida 
and Texas. It also includes North American fossils, as well as a num- 
ber of Recent species from southeast Asia (Korea, China, and Thailand). 
All the species called "Pleurocera" by Bryant Walker, and other 
authors who followed him blindly, and all the species called "Goni- 
obasis" (with very few exceptions) belong to this genus. Their eggs 
are laid in a single row in a close, irregularly spiral group, in apparent 
flat clusters of 3 to 10 egg capsules in each small egg mass, the whole 
covered with sand grains as recorded by Van Cleave (1932), Winsor 
(1933), and Woodward (1934). 



360 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Rafinesque originally described Oxytrema (1819, p. 423), including 
young specimens of both, his genus Pleurocera and this one. There 
were no specific names included, Blainville, in 1824 and again in 
1825 (p. 442), placed one species of Rafinesque under this generic 
name. He gave as the sole example of the subgenus Pleurocerus 
(Oxytrema) acutus (Rafinesque), validating the specific name in 1824. 
Rafinesque (1831, p. 3) again described his Pleurocera acuta, and at 
the same time declared he had given the name in 1818. From 1824 
on, Oxytrema has been the earliest available name for the group be- 
cause the genotype was fixed at that time as Pleurocera (Oxytrema) 
acuta Blainville by monotypy. With no serious question ever raised 
about the identity of acuta, the genotype, doubts about the identity 
of the genus Oxytrema vanish. 

The eastern American (Appalachian) species of Oxytrema include 
some whose shells are almost completely smooth as adults, such as 
0. symmetrica (Haldeman) ; some with spiral sculpture predominant, 
such as 0. drginica multilineata (Say); others with axial sculpture 
strong, such as 0. laqueata (Say); and still others with both axial 
and spiral lirae to produce reticulate or nodose sculpture, such as 
0. catenana (Say). 

In exactly parallel fashion, as should be expected of congeneric 
stocfe, the western American species now laiown to belong to Oxytrema 
show the same rather complete range of sculpture from smooth to 
axially and spu-ally sculptured adult shells. The easternmost living 
representative of the Rocky Mountain group of species is 0. comalensis 
(Pilsbry). This species from Texas belongs to this minor group 
within the genus, possessing the same minutae of female reproductive 
characters as does 0. plicifera (Lea) from Oregon and Washington. 
Most if not all the western America Tertiary fossil species described 
as "Melania," "Goniobasis," and ''Pachychilus," from Texas to 
Washington, were members of this genus Oxytrema Rafinesque. For 
example, ''Amhloxus'' olequaensis Arnold and Haiuiibal (Hannibal, 
1912, p. 178, pi. 8, fig. 27) closely resembles the smoother phase of 
the living Oxytrema silicula (Gould) from the same region, while 
"Pachychilus" drakei Arnold and Hannibal (Hannibal, 1912, p. 183, 
pi. 8, fig. 26) is close to the living species Oxytrema plicifera (Lea). 
Certain other fossil species from the Rocky Mountain region parallel 
eastern forms in the possession of cancellate or reticulate sculpture. 
Species known from the fossil record indicate the previous continuity 
of geographic distribution from Washington to Texas, now noncon- 
tinuous since much of the intervening area (progressively desiccated 
since the Miocene) is now unsuited to survival of pleurocerine fresh- 
water snails. 

Female animals of Oxytrema canaliculatum undulatum (Say) from 



OLD AND NEW WORLD MELANIANS — MORRISON 361 

the Kentucky River, of Oxytrema deshayesiana (Lea) from the Tennes- 
see River, of Oxytrema bvlbosa (Gould) from eastern Oregon, and of 
Oxytrema nodifila (Martens) from the Han River, Korea, have been 
examined by me and sketched to show the egg-laying groove and the 
"ovipositor" pit on the right side of the foot (see pi. 11). 

Section Strephobasis Lea, 1861 

The group named Strephobasis by Lea (1861, p. 96), with the geno- 
type Oxytrema (Strephobasis) plena Anthony, as stated by Pilsbry 
(1896a, p. 496), is only an extreme section of the genus, according to 
shell characters. This small group, which is confined to the larger 
rivers of the upper Tennessee drainage, is exactly parallel to Goniobasis, 
sensu stricto, from the Coosa River di'ainage. The animal characters 
must be examined to determine whether this group is identical to or 
biologically distinct from Oxytrema, sensu stricto, and Goniobasis, 
sensu stricto. 

Genus Gyrotoma Shuttleworth, 1845 

Gyrotoma Shuttleworth (1845, p. 88), based on the genotype 
(Gyrotoma ovoideum Shuttleworth =) Gyrotoma excisum (Lea), is con- 
fined to the Coosa River drainage. As Goodrich (1924, p. 6) has 
hinted, the group named Goniobasis by Lea in 1862 (genotype by 
subsequent designation by Hannibal 1912: G. osculata Lea) may 
eventually prove to be most closely related to Gyrotoma. It is possible 
that Goniobasis, sensu stricto, may be a subgenus of Gyrotoma different 
only by lack of or development of the fissure in the lip. Study of the 
animal characters (as yet unknown) of the genotype must prove the 
true biological position of Goniobasis. 

Genus Mudalia Haldeman, 1840 

Mudalia of Haldeman, whose genotype is Mudalia carinata (Bru- 
guiere) from the Atlantic coastal region, includes trilineata (Say) of the 
Ohio River and the more widespread "Goniobasis" livescens (Menl^e) 
from the upper St. Lawrence River and upper Mississippi River areas. 
The single carina typical of adolescent shells and the egg-laying habits 
are identical in carinate (Winsor, 1933) and in livescens (Jewell, 1931). 
Figures 10 and 14 of Goodrich (1945, pi. 1) show how closely the shape 
of adult shells of livescens may vary toward the usual shape of adult 
shells of carinata. The eggs of Mudalia are laid singly; that is, there is 
only one egg capsule in each egg mass. 

Genus Leptoxis Rafinesque, 1819 

Leptoxis of Rafinesque is the earliest name available for the Ameri- 
can species related to the genotype L. praerosa (Say), and must be so 
used. Pilsbry (1917, p. 113) has stated the case for priority of Lep- 



362 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

toxis Rafinesque 1819 over Anculosa Say 1821 so clearly that Walker's 
1918 refusal to follow this needed correction of nomenclature appears 
at present to be only a prejudiced personal objection to the change. 
As in other similar cases, most American authors, especially those 
concerned with biology rather than nomenclature, from 1918 to date 
have followed Walker not because they had critically checked the 
generic names used by him, but simply because it was the most recent 
comprehensive classification outline for North American fresh-water 
shells. As stated elsewhere, this incorrect usage following Walker 
dates only from Walker, and has only 33 years standing, which is not 
enough to even raise any questions of usage over priority of names. 

For the record Chenu states (Haldeman, 1847-1848, p. 1, footnote) 
that he and not Haldeman put the name Leptoxis of Rafinesque on 
Haldeman's pictorial monograph of the genus. Thus Chenu, and 
Pilsbry by formally designating praerosa as genotype in 1917, com- 
pletely cleared up all doubts, concerning Rafinesque's genus, which had 
existed prior to 1848. 

Genus Eurycaelon Lea, 1864 

Eurycaelon of Lea (1864, p. 3) has as genotype Eurycaelon anthonyi 
(Redfield) by subsequent designation by Walker (1918, p. 36). The 
two known species, Eurycaelon crassa (Haldeman) 1841 and E. 
anthonyi (Redfield) 1854 are confined to the larger rivers of the upper 
Tennessee drainage as reported by Goodrich (1931). In this con- 
nection must be mentioned the fact that the two names cristata 
Anthony (young specimens) and anthonyi Redfield (adult specimens) 
were published simultaneously in April 1854, according to the printed 
signature dates of volume 6 of the Annals of the Lyceum of Natural 
History of New York. Goodrich, acting as first reviser, selected 
anthonyi Redfield, on page 130 of volume 6 of the above Annals, over 
the name cristata of Anthony, on page 108, which he placed in syn- 
onymy. In this way Goodrich avoided a confusing change of name 
of the genotype. 

Genus Pleurocera Rafinesque, 1818 

Pleurocera of Rafinesque became monotypic in 1820 with the valid 
publication by Rafinesque of Pleurocera verucosa. Try on in 1864 and 
1873 did not include the genotype in his usage, so his usage was bio- 
logically and nomenclatorially wrong. Pilsbry in 1896 questioned the 
genotype. Hannibal (1912, p. 169) formally designated the mono- 
type species verrucosa Rafinesque as the genotype, being followed cor- 
rectly by Pilsbry in 1917. 

Bryant Walker, in his 1918 classification, continued to use Tryon's 
incorrect name for the genus when he used only the second available 



OLD AND NEW WORLD MELANLANS — MORRISON 363 

name, Lithasia Haldeman (1840), which has the species geniculata of 
Haldeman for genotype. 

The smooth species from the Kentucky River named Ellipstoma 
zonalis by Rafinesque in 1818 can only be the species commonly 
known as Lithasia obovata Say 1829. With over 10 years priority, the 
specific name of Rafinesque must be used. Animals of Pleurocera 
zonalis Rafinesque personally collected from the Kentucky River 
have been examined and sketched. The female egg-laying characters 
are illustrated in figure 4 of plate 11. 

Subgenus Ellipstoma Rafinesque, 1818 

In spite of the extreme brevity of Rafinesque's original description 
(1818b, p. 42), critical reading indicates that this name applies only 
to the gi'oup afterward named Angitrema by Haldeman (1841a). 
Hannibal (1912, p. 168) formally designated Rafinesque's first species, 
Ellipstoma gibbosa, as the genotype, although he misidentified that 
species. When one approaches the identification of gibbosa from 
every angle it is evident that there is only one kind of snail in the 
Ohio and Wabash Rivers with "a large knob behind the outward lip." 
This is the same species namedj almost^three years later as Melania 
armigera Say (1821, p. 178). 

Critical examination of the animal characters of the genotype 
Pleurocera (Ellipstoma) gibbosa Rafinesque is needed to check the bio- 
logical distinction of Ellipstoma as a subgenus, a separate genus, or a 
synonym of Pleurocera, sensu stricto. 

Genus Anaplocamus Dall, 1895 

Anaplocamus of Dall (1895, p. 8), mistakenly described as an Alaskan 
marine shell because of accidentally transposed locality labels, is a 
monotypic genus based on the species "Anculosa" dilatata Conrad 
(Rehder, 1942, p. 49). This species of the Kanawha River drainage 
and the upper Cheat River of the Allegheny drainage is the northern 
representative of the lo group. It has the columellar shell structure 
of lo, not that of Leptoxis. From a broad biological viewpoint it 
may be considered a subgenus of lo, or, more conveniently, a separate 
genus closely related thereto. 

Genus lo Lea, 1831 

The genus lo of Lea is well known; it includes the largest living 
United States Pleuroceridae known. How many species other than 
the nomiual genotype fluvialis (Say) there are or were in existence 
must await studies of the animals to check the magnificent analyses 
of shells by Adams (1915). Haldeman, in the American edition of 



364 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loa 

Heck's Iconographic Encyclopaedia (1851, p. 84), has given a general 
description of the animal of lo : 

Its characters and habits are not those of Fusus, but of [Oxytrema] proper, as 
distinguished from Leptoxis; for although it inhabits the rapids as well as quiet 
water, in both cases it avoids the current by seeking shelter beneath shelving 
rocks, or in hollows or crevices in them. The head is large, and with the tentacles 
much exposed; the foot is^as large^as inr [Oxyirema]; the -coloration is the same 
(black lines upon an orange ground) ; the operculum is subspiral as in [Oxytrema], 
the mantle extends into the canal of the shell, but does not form a closed siphon; 
the vent is upon the right side; the mouth is a longitudinal slit; the eyes (which 
are sensitive to the light) are upon a short enlargement of the outer base of the 
tentacles, which may be a little longer than in [Oxytrema], and they are not visibly 
annulated. It differs from Fusus, and resembles [Oxytrema] in living upon vege- 
table food; and it moves along in a sluggish manner, moving the head from side 
to side upon the bottom. lo spinosa and I. tenebrosa are merely varieties of /. 
fluvialis. The spinose individuals are much the most abundant, although the 
species is rare when compared with various species of [Oxytrema]. 

We have taken the liberty of replacing the name "Melania" in the 
foregoing quotation with that of Oxytrema, since Haldeman had 
studied and published as typical of "Melania" the animal of Oxytrema 
virginica (Gmelin) (Haldeman, 1841b, p. 21). 

In this connection, it must be mentioned that the shells of the group 
of " Pleurocera" alveare Conrad (1834, p. 54, pi. 4, fig. 7) look sus- 
piciously like normal sized representatives west of and downstream 
from the megasomatic lo, sensu stricto, species. The generic name 
Megara H. and A. Adams (1854, p. 306) has not hitherto been geno- 
typed. Walker (in litt.), quoted by Goodrich (1928, p. 2, footnote 5), 
incorrectly reported that Hannibal (1912, pp. 169, 179) had designated 
the type of Megara. Reference to these pages shows that Hannibal 
listed parts of the original Megara twice, with two different species as 
examples (not as types) in the two synonymy lists 10 pages apart. 
The genotype here designated is the first species, Megara alveare 
(Conrad) 1834. Animal characters of this group remain a desideratum. 
If the group is most closely related to Oxytrema, as it has usually been 
regarded, Megara will remain as a synonym of Oxytrema. If, however, 
these species should prove upon examination to be closest to lo in 
relationship of animal characters, their group name Megara will be 
available. 

Subfamily pleurocerinae (in Central and South America) 

Genus Pachychilus Lea, 1851 

This genus seems to be the ecological replacement of Oxytrema, to 
the southward of the United States. In my opinion, all the known 
species of this family and subfamily from Central America and the 
West Indies belong to Pachychilus. In this connection, the ichthyol- 



OLD AND NEW WORLD MELANUNS — MORRISON 365 

ogist S. E. Meek should be given credit as the first known discoverer 
of the eggs of any member of the family Plem-oceridae. Reporting on 
collections made in 1906 (Meek, 1908, pp. 205-206), he says: 

Sphaeromelania largillierti Phil. This is the largest and by far the most abundant 
species of shell found in the lake. It is found everywhere along the shore on plants 
and rocks. This species deposits its eggs in January and February. The eggs are 
very large and are usually attached to the under side of rocks. This species is so 
abundant that its large gelatin-like eggs would furnish a considerable amount of 
food for small fishes. It is a very abundant species in Guatemala and San Salvador 
to Central Nicaragua. 

Goodrich and Van der Schalie (1937, pp. 39, 41, 42) have recorded and 
described the eggs of three additional species of Pachychilus as follows: 

Pachychilus glaphyrus (Morelet) 1849. The eggs were found attached to dead 
leaves, which were abundant in the bottom of the arroyo. The leaves were in 
only a few inches of water on a mud-silt bottom near shore. The eggs occur in 
groups of 3 to 5, are round and glassy in appearance, and are irregularly and line- 
arly arranged as a single loosely attached layer. As might be expected in a 
species as large as glaphyrus, the eggs are also large, having a diameter of 3.9 mm. 
(Arroyo Yalchactila of the Rio de la Pasion, about 4 mi. S. W. of La Ceiba, Alta 
Vera Paz, Guatemala) .... 

Pachychilus corvinus (Morelet) 1849. Egg masses of this species were found 
from which the infant shells were emerging. These were smooth, pinkish carnelian, 
translucent, and bluntly rounded at the periphery. The whorls varied from 3 to 
3H- With growth the shell generally becomes black, even though the adults may 
be of a mahogany or yellowish color on the body whorls .... 

Pachychilus pilsbryi Martens 1897. Eggs of this species were found on the lower 
surfaces of stones in a current near shore. The egg masses occur as flat layers 
spread out over flat objects. The size of the egg mass varies, the number of eggs 
in some of them observed running from 8 to 20. The eggs themselves are round, 
shotlike, and transparent. The whole mass is held together by a gelatinous 
coating. The average size of the individual eggs is 2.8 mm. 

We are indeed fortmiate in having on record, in the first four species 
so known, the egg-laying characters of three subgenera of the genus 
Pachychilus. 

Subgenus Oxymelania Crosse and Fischer, 1892 

Oxymelania Crosse and Fischer (1892, p. 328) possesses the species 
schiedianus Philippi as genotype by original designation. This group 
of small to medium-sized species has been most completely studied 
in the San Luis Potosi region by Pilsbry, following the collections 
made by A. A. Hinkley. In this subgenus, some species of which are 
widely different in height of spire, and thus of shell outline, the single 
general identifying character is the possession or prominence of macro- 
scopic spiral lirations on the adult shell. The egg-laying habits for 
the species pilsbryi Martens are quoted above. The synonyms of 
Oxymelania include Potamanax Pilsbry (1893, p. 340), named only a 
few months later, and Lithasiopsis Pilsbry (1910, p. 47). In spite of 

273553—54 2 



366 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loa 

the great similarity of shells, Guhaedomus does not belong here. Ani- 
mal characters prove it to belong in the family Thiaridae, as recorded 
below. Oxymelania is known to range geographically from the Panuco 
River system of Mexico to the headwaters of the Usumasinta River 
in Guatemala. 

Subgenus Pachychilus Lea, sensu stricto, 1851 

Pachychilus Lea (Lea and Lea, 1851, p. 179) originally included 
only two specific names, cumingii Lea and laevissima Sowerby. In 
January 1860, Reeve (1859-1861, pi. 18, fig. 126) designated laevissima 
as the genotype. On the other hand, Reeve's "laevissima" was a 
mixture of the species from Chiapas, Mexico, which has since been 
mistakenly called "laevissima" by many authors, and the true laevis- 
sima of Sowerby and Lea from La Guayra, Venezuela. Since the 
mixture of Reeve contained in synonymy the originally included 
laevissima of Sowerby, this designation was valid. Thus the genotype, 
correctly figured on plate 19, figure 133c, of Reeve (1859-1861) is 
the only known South American species of the genus. The typical 
group in Central America is the corvinus or largillierti group of species, 
characterized by medium- to large-sized shells with a generally elongate- 
ovate or elongate-conic outline, rather flat-sided whorls, and almost 
completely smooth macroscopic sculpture. The egg-laying habits are 
quoted above. The Cuban species nigrata Poey 1858, fuentesi Aguayo 
1936, and violaceus Preston 1911 apparently belong here. Synonyms 
of Pachychilus, sensu stricto, include Cercimelania Crosse and Fischer 
(1892, p. 327) with P. liehmanni (Philippi) as genotype, and Sphaero- 
melania Rovereto (1899, p. 109) with the same genotype as Pachy- 
chilus, since it was proposed as a substitute name. Apparently both 
Crosse and Fischer and Hannibal did not know of Reeve's type desig- 
nation and incorrectly listed P. graphium (Morelet) as the genotype. 

Subgenus Glyptomelania Crosse and Fischer, 1892 

Glyptomelania Crosse and Fischer (1892, p. 328) is confined to 
Central America. This group of large shells with variable to nodose or 
subspinose sculpture includes glaphyrus (Morelet), the subgenotype 
by original designation. The eggs of this species have been recorded 
by Goodrich and Van der Schalie as quoted above. 

Subgenus Pilsbrychilus Morrison, 1952 

Pilshrychilus Morrison (1952, p. 7) is distinguished from other known 
members of the genus by the markedly sinuous lip-margin developed 
in mature adult shells. It seems probable that examination of the 
animal will serve to corroborate the distinction of this group now based 



OLD AND NEW WORLD MELANIANS — MORRISON 367 

solely on shell characters. The genotype by original designation is 
Pachychilus {Pilsbrychihs) dalli Pilsbry, which was first validly 
published April 17, 1896, in Science (vol. 3, p. 608) ahead of the com- 
plete description by Pilsbry in Proceedings of the Academy of Natural 
Sciences of Philadelphia, June 16, 1896 (p. 269). Fortunately, this 
earliest validation preceded the typographical error "walli," which 
appeared in Zoologischer Anzeiger in May 1896. 

Genus Doryssa H. and A. Adams, 1854 

Doryssa H. and A. Adams (1854, p. 304), whose genotype is D. 
atra (Bruguiere) from Suriname, ecologically and geographically 
replaces Pachychilus in northern South America from Venezuela to 
the Guianas and Brazil. In both these genera the females have a 
well-developed genital groove ending in the characteristic pleurocerine 
glandular egg-laying sinus near the right edge of the foot. The 
males lack this groove. Doryssa consolidata (Bruguiere) was per- 
sonally collected by the hundreds from rocks in the Mazaruni and 
Cuyuni Rivers of the Essiquibo system, British Guiana, in the 
summer of 1925. At that time the shells were cleaned and the 
radulae (see Baker, 1930, p. 30) extracted from the animals for future 
study of variation. Although at that time I did not know what char- 
acters were critical for subfamily classification, no difference of gross 
anatomy was observed between this British Guiana Doryssa and the 
North American species Oxytrema semicarinata (Say), already crit- 
ically and completely personally studied near Lexington, Ky., in 
1924. The egg-laying season of Doryssa is apparently still unrecorded. 
It was not observed during June, July, or August in the Kartabo 
region. Sheppardiconcha Marshall and Bowles (1932, p. 3), described 
as a new genus from the (Miocene?) fossil deposits of the Upper 
Amazon region of Ecuador, is not a subgenus of Aylacostoma. Refer- 
ence of the group by later authors to Aylacostoma (=Hemisinus) is 
primarily due to the difficulty of complete separation of the genus 
Doryssa of the Pleuroceridae from Aylacostoma of the Thiaridae by 
shell characters alone. According to the shell characters of the 
original material, Sheppardiconcha is a synonym of Doryssa, 

Family Pleuroceridae (in Asia) 

Subfamily pleurocerinae 

Genus Oxytrema Rafinesque, 1819 

In the present state of our knowledge it is no longer possible to 
maintain the generic distinction of certain eastern Asiatic species 
from the genus Oxytrema, which has previously been considered 



368 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

wholly North American. The generic name Hua, proposed by S. F. 
Chen (1943, p. 21) for the group of "Melania" telonaria Heude 1888, 
is a synonym of Oxytrema Eafinesque 1819. As is the case in both 
the Appalachian and Rocky Mountain regions of North America, 
the genus Oxytrema is represented in eastern Asia (Korea, China, 
and Thailand) by some species that are generally smooth (Hua) 
and some with strongly nodose or reticulate sculpture. Heude 
figured the female pleurocerine external anatomy of Oxytrema in 
1890 in his excellent work on the Chinese fresh-water mollusks. 
Unfortunately, he misunderstood the structures of the females of 
such species as "Melania" jacquetiana Heude and called them male 
individuals. At present, Heude's mistake is easy to explain. The 
(female) egg-laying apparatus of these species (Heude 1890, pi. 43, 
fig. 5) is superficially identical in appearance to the (male) intromittent 
organ of certain operculate land snails of the subfamily Cyclophorinae 
(family Cyclophoridae) (see Heude 1890, pi. 42, fig. 12c), as a com- 
parison of Heude's own figures will demonstrate. It is this extreme 
similarity in appearance that led Heude to an understandable mis- 
interpretation and has effectively hidden the complete understanding 
of his critical discoveries for many years. 

The present studies have corroborated those of Heude. The 
animals of Oxytrema telonaria (Heude) 1888 are dioecious; the sexes 
may be distinguished by the presence or absence of the female genital 
groove on the right side of the foot. There is a glandular, egg-laying 
pit at the terminal end of the groove a little above the right edge of 
the foot. The epithelium lining this groove is markedly different 
from that of the adjacent area and is easily distinguished in preserved 
material. The posterior edge of the groove is projected into the pit 
as a sort of papilla at its termination. The oviduct is not enlarged 
into a uterus; there are no embryos in the body behind the head. 
Animals of the followmg Asiatic species have been personally examined 
and found to exhibit the same pleurocerine reproductive anatomy 
with but little difference in minor details: 0. nodijila (Martens) 1886, 
Han River, Korea (see pi. 11, fig. 3); 0. toucheana (Heude) 1888, 
Min River, Fukien, China; 0. jacquetiana (Heude) 1890, Shaohsing, 
Chekiang, China; 0. peregrinorum (Heude) 1890, Fukien, China; 
0. moutoniana (Heude) 1890, Sa-Hsien, Fukien, China; 0. joretiana 
(Heude) 1890, Chekiang, China; and 0. bailleti (Bavay and Dautzen- 
berg) 1910, Szechwan, China. How many of the other Asiatic species 
listed by S. F. Chen in these groups will prove to belong to Oxytrema 
can be determined only when their reproductive anatomy is examined. 
The presence of Oxytrema species in the recent fauna on both sides of 
the North Pacific Ocean demonstrates a geographic distribution similar 
to that of the genera Viviparus and Anodonta, sensu stricto. It is 



OLD AND NEW WORLD MELANIANS — ^MORRISON 369 

evident also that the western American (Alaskan) region consti- 
tuted the pathway of migration of these fresh-water mollusks between 
Asia and North America. 

Genus Paludomus Swainson, 1840 

With a short, ovate shell-shape corresponding to that of the Ameri- 
can genus Leptoxis, the Asiatic genus Paludomus (Swainson, 1840, 
pp. 198, 340) also belongs to the subfamily Pleurocerinae. The 
anatomy of the animals of Paludomus tanschaurica (Gmelin) has 
been completely studied by Seshaiya (1934). The finding by Seshaiya 
of a tubular structure which he called a "penis" along the terminal 
portion of the primary gonoduct of the male must be discounted as 
a glandular discovery until it is proven to be an intromittent organ. 
As Seshaiya himself pointed out, it is not muscular as is the case 
of the similarly located and functional male organ of the Tiphobiinae. 
Seshaiya believed these animals oviparous, as he found no evidence of 
ovoviviparity in their anatomy. He did not, however, mention either 
the egg-laying groove or the pit. Paludomus labiosus (Benson) (?) 
from Thailand and Paludomus maculatus Lea from India (also 
members of the subgenus Paludomus, sensu stricto, with its concentric 
operculum) have been personally examined. The female animals are 
pleurocerine, with an egg-laying groove and pit in the side of the foot 
and without any brood pouch (see pi. 11, fig. 6). The true biological 
relations of the other subgenera such as Hemimitra, which differ 
markedly in shell and opercular characters, are still unknown or 
unrecorded. 

Subfamily lavigeriinae Thiele, 1929 

Members of the subfamily Lavigeriinae Thiele (1929, p. 79) show 
the full dioecious mode of reproduction of the family. That is, there 
is a normal 1:1 sex ratio of males to females as in all other known 
members of the Pleuroceridae. The most important distinguishing 
character is the possession of a uterine brood pouch similar to that in 
the Viviparidae. As in the typical subfamily (Pleurocerinae), the 
males possess no intromittent structures whatsoever. Coincident to 
the development of ovoviviparity, the females have lost (at least do 
not possess) the egg-laying sinus and papilla characteristic of the 
Pleurocerinae. The name Semisulcospirinae Morrison (1952, p. 8) is 
a synonym of the earUer subfamily name Lavigeriinae. 

Genus Semisulcospira Boettger, 1886 

The genotype of Semisulcospira Boettger (1886, p. 4) is the well- 
Icnown Japanese species Semisulcospira libertina (Gould) 1862. Ex- 
amination of many specimens from the Hayakawa^River, ^Honshu, 

273553—54 3 



370 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

Japan, sent to the U. S, National Museum a few years ago by Captain 
Honess, U. S. Army Sanitary Corps, conclusively demonstrated that 
this species is ovoviviparous, as previously suspected. The head and 
snout of these snails are uncomplicated in either sex by external re- 
productive structures of any kind. Coitus is probably accomplished 
by apposition of the mantle edges, with perhaps some prolapsis of the 
genital duct openings. The lower part of the female reproductive 
duct (oviduct) is enlarged to form a functional uterus. It is thin- 
walled when fully distended with young and apparently is not glandu- 
lar. The hundreds of embryonic young of uniformly small size 
developed at one time appear to be packed in axial rows, visible ex- 
ternally when the animal is removed from the shell and resembling 
the conglutinates of glochidia of the North American fresh-water 
mussel genus Strophitus. In some cases the young nearest the ovi- 
duct are not fuUy developed, showing that development of the ova is 
at least partly progressive as the young approach the uterine mouth. 
On the other hand, the great uniformity of size of all the fully de- 
veloped young, with shells of about 2 whorls, would indicate a definite 
seasonal, mass release of the young. At any rate, there is no indica- 
tion of any continued growth to large size of a few embryos at a time 
or continued parturition over a long season or throughout the year 
as in certain genera of the family Thiaridae. 

Kecently, Corp. W. E. Old of the U. S. Army Signal Corps sent 
four species of "Alelania" from Korea to the U. S. National Museum. 
Examination of the animals by both R. T. Abbott and the author has 
proven that "Melania" nodijila Martens is an Oxytrema as noted above, 
while the species gottschei Martens, nodiperda Martens, and graniperda 
Martens (1905) are members of the genus Semisulcospira. The 
geographic range of Semisulcospira across continental Asia is imper- 
fectly known and in need of continued research. 

Family Pleuroceridae (in Africa) 

Subfamily pleurocerinae 

Genus Potadoma Swainson, 1840 

The genotype of Potadoma Swainson (1840, pp. 200, 341), P.freethii 
(Gray), must be examined to completely clear up the position of this 
genus. Pilsbry and Bequaert (1927) have reported that these snails 
are apparently oviparous, with the edge of the mantle smooth and even. 
They compare these species to some species of the American genus 
Pachychilus. If Potadoma is oviparous as reported, with the egg-laying 
groove, it belongs in the subfamily Pleurocerinae, and the subfamilies 



OLD AND NEW WORLD MELANIANS — MORRISON 371 

Potadominae Pilsbry and Bequaert and the Paludominae are included 
synonyms. In addition to the species called Potadoma by Pilsbry 
and Bequaert, some of the species they called ''Melanoides" also belong 
to Potadoma of Swainson. "Melanoides" species such as ignohilis 
(Thiele) (Pilsbry and Bequaert, 1927, p. 278, pi. 26, figs. 1, 2) are 
smooth; others such as medjeorum Pilsbry and Bequaert (1927, p. 283, 
fig. 47) and tornata (Martens) (Pilsbry and Bequaert, 1927, p. 283, fig. 
48) possess strong spiral sculpture. Still others, such as wagenia Pilsbry 
and Bequaert (1927, p. 262, pi. 22, figs. 14-16), crawshayi (E. A. Smith) 
and mweruensis (E. A. Smith) (Pilsbry and Bequaert, 1927, pp. 264- 
265, pi. 22, figs. 19-22) show strong plicate to cancellate shell sculpture. 
The total picture of sculpture in this genus seems exactly parallel to 
that seen in Appalachia, western America, and eastern Asia in the 
genus Oxytrema. 

Genus Limnotrochus E. A. Smith, 1880 

1880. Limnotrochus E. A. Smith, 1880b, p. 425. (Genotype, Limnotrochus 

thomsoni, E. A. Smith, 1880.) 

Genus Paramelania E. A. Smith, 1881 

1881. Paramelania E. A. Smith, 1881b, p. 559. (Genotype, Paramelania 

damoni E. A. Smith, 1881, by subsequent designation by Pilsbry and 
Bequaert, 1927, p. 320.) 

Genus Spekia Bourguignat, 1879 

1879. Spekia Bourguignat, 1879, p. 27. (Genotype, Spekia zonatus (Woodward), 
1859, by monotypy.) 

Pilsbry and Bequaert have said that, as far as known, the three 
above-named genera are oviparous. If complete study of the animals 
confirm-s this and reveals the characteristic egg-la3ang pit in the side 
of the foot, these Lake Tanganyika forms are also Pleurocerinae. 
Moore (1899a, p. 171) found neither an external groove nor a brood 
pouch of any type in the females of Spekia zonafa (Woodward) 
examxined by him. This condition seems identical in appearance to 
that of those North American members of the pleurocerine genus 
Oxytrema, in which the female egg-laying groove is not demarcated 
structurally and can only be seen functionally as a linear depression 
in the skin during the breeding season. 

Subfamily lavigeriinae Thiele, 1929 

The statement of Pilsbry and Bequaert (1927, p. 300) in regard to 
the ovoviviparity of the mixture of species they called "Melanoides" is 
in need of clarification. They left the problem unsolved when they 
said: "Typhohia, Bathanalia, and Lavigeria are viviparous, having the 



372 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

last part of the oviduct enlarged to form a brood pouch in the mantle, 
as in Melanoides and Viviparus." The fact is that Melanoides does 
not have a uterine brood pouch as in the family Viviparidae. True 
Melanoides species have the subhaemocoelic brood pouch in the back 
of the neck, with the brood pouch pore on the right, as in all other 
members of the Thiaridae. 

If Pilsbry and Bequaert actually saw uterine brood pouches in the 
animals of any African species of "Melanoides," they probably were 
dealing with members of the subfamily Lavigeriinae, of an unnamed 
genus. In this connection, the possibility should be stated frankly 
that the subfamilies Lavigeriinae and Tiphobiinae may in the future 
prove to be biologically confluent. Much more study of the animals of 
African fresh-water species both in and out of Lake Tanganyika is 
needed to prove or disprove this possibility. As defined above, the 
subfamily Lavigeriinae includes those genera of pleurocerid snails in 
which the males have no intromittent organ and the females possess 
uterine brood pouches. 

Genus Lavigeria Bourguignat, 1888 

The genus Lavigeria Bourguignat (1888, p. 33) is another of the 
peculiar pleurocerid forms found in Lake Tanganyika. In addition to 
the shell characters, this genus differs from others by its operculum, 
which is modified by partial uncoiling to a subspiral form. Moore 
1899b, p. 192, pi. 20, fig. 6) has furnished anatomical studies of the 
species Lavigeria coronata Bourguignat. He called the animal "Nas- 
sopsis nassa Woodward," while Pilsbry and Bequaert have referred 
this anatomical study to the genotype which they designated, namely: 
Lavigeria grandis (E. A. Smith) 1881. Moore's figure of the shell 
proves he studied the species coronata. He found that this genus 
possesses a uterine brood pouch similar to that of Viviparus and of 
the Tiphobiinae. According to him, the males do not possess any 
secondarily developed intromittent organ such as is found in the 
mantle edge of the males of Tiphohia. 

Genus Bourguignatia Giraud, 1885 

1885. Bourguignatia Giraud, 1885, p. 193, pi. 7, figs. 5-7. (Genotype, Bourguigna- 
tia imperialis Giraud, 1885, by monotypy.) 

Genus Edgaria Bourguignat, 1888 

1888. Edgaria Bourguignat, 1888, p. 33. (Genotype, Edgaria paucicostata (E. A. 
Smith), 1881, by subsequent designation by Pilsbry and Bequaert, 1927, 
p. 328.) 

The shells of the Bourguignatia, Lavigeria, Edgaria, and Paramelania 
complex of Lake Tanganyika are almost identical in general characters ; 
however, there are two distinct types of opercula known in the group. 



OLD AND NEW WORLD MELANL/USTS — MORRISON 373 

Our present lack of anatomical knowledge of all but Lavigeria makes 
it impossible to say whether two or more subfamilies are represented 
in this complex of shells inhabiting the rocky shorelines of Lake 
Tanganyika. 

Genus Bythoceras Moore, 1898 

1898. Bythoceras Moore, 1898c, p. 452. (Genotype, Bythoceras iridescens Moore, 
1898, by monotypy.) 

These shells from deeper waters of Lake Tanganyika are also very 
similar to those of the Lavigeria complex; their opercula differ from 
those of Lavigeria in being secondarily concentric around a paucispiral 
nucleus. The general statement by Moore (1898d) that Bythoceras 
is much more like Tanganyicia in anatomy than (" Nassopsis"==) 
Lavigeria is in need of corroboration or clarification. Neither 
is it clear from Moore's subsequent statements and figures (1899b) 
whether Bythoceras is ovoviviparous or not. 

The final allocation of these names and also of Paramelania and the 
subfamily name Paramelaniinae must await the complete recording 
of shell, opercular, and reproductive characters of all members of the 
Lavigeria complex of Lake Tanganyika. If the reproductive anatomy 
proves identical, Paramelaniinae (Moore, 1898b, p. 315) will super- 
sede the name Lavigeriinae (Thiele, 1929, pp. 79, 83) used at this time. 

Subfamily tiphobiinae Moore, 1898 

These are pleurocerid snails modified in the males by the develop- 
ment of a secondary, eversible "penis" or vergic structure in the 
mantle edge near the end of the vas deferens. As far as known, these 
are the only "Melanian" snails of any kind that possess any intro- 
mittent structures whatsoever. The females are ovoviviparous, with 
the terminal portion of the oviduct modified into a uterine brood 
pouch for the young. At present, our knowledge — or lack of 
knowledge — indicates that the Tiphobiinae are solely African in 
geographic distribution, parallel to but not identical with the Asiatic 
and African subfamily Lavigeriinae. 

Genus Tiphohia E. A. Smith, 1880 

The genotype of Tiphobia E. A. Smith (1880a, p. 348, pi. 31, fig. 6), 
T. horei E. A. Smith, 1880, possesses the reproductive characters of 
the subfamily, as defined above, according to the studies of Moore 
(1898a). The statement of Pilsbry and Bequaert that the characters 
of the Tiphobiidae do not distinguish them from the Melaniidae is 
based on their misunderstanding of the true nature of the brood 
pouch of the Thiaridae ("Melaniidae"). Tiphohia is one of the 
endemic genera of Lake Tanganyika, with a spinose shell and a 



374 PROCEEDINGS OF THE NATIONAL MUSEUM vol. los 

secondarily concentric operculum. Its complete biological relation- 
ship with the other Tanganyika genera and with extra-Tanganyika 
forms must be clarified by future studies. 

Genus Bathanalia Moore, 1898 

The genotype, Bathanalia howesi Moore, 1898, is stated by Moore 
(1898a) to be "almost identical anatomically with Tiphobia horei." 
This genus differs from Tiphobia by shell characters such as height of 
spire and the perforate columella. In addition, Bathanalia possesses 
an unmodified (paucispiral) operculum. 

Family Thiaridae 

The pattern of reproduction is the same for all known members of 
the family Thiaridae. A high degree of specialization in this one 
character is indicated by this observed pattern of ovoviviparous 
parthenogenicity. In other words, there is no male indiv^idual known 
in this entire family of snails. To my knowledge, the parthenogenesis 
of the Thiaridae has not been examined cytologically to determine 
whether it is haploid or diploid. 

There is required only one individual (any individual) to start 
a new population or colony if it reaches a new fresh-water locality by 
stream capture or by adventitious transportation. The resultant 
ability of these snails theoretically to spread more rapidly may partly 
explain their wider distribution in suitable habitats on islands in both 
the Pacific and the West Indies and their wider expansion southward 
across South America than that exhibited at present by the family 
Pleuroceridae. Certain corollaries are assumed in this theoretical 
explanation of the differences and peculiarities of distribution of the 
two fresh-water families of Pleuroceridae and Thiaridae. Structurally, 
the animals of these families are relativel}^ primitive, exceedingly so 
as regards their fundamental reproductive characters. Without any 
proof to the contrary, we may assume them to be of approximately 
equal antiquity, geologically speaking. Reasoning from the special- 
ized pattern of reproduction of the Thiaridae, we may logically argue 
that they are the younger group evolved from the older ancestral type 
represented now in fresh water by the Pleuroceridae. If this be true, 
the younger has outstripped the more primitive older type in geo- 
graphic expansion over geologic time from the era of their differentia- 
tion up to the present. If the Thiaridae were ever in North America 
we must assume that they are no longer represented there because the 
conditions of existence are not now or were not, during some previous 
geologic era, favorable to their continued existence in those northern 
areas. Their expansion across the North American continent may be 



OLD AND NEW WORLD MELANIANS — MORRISON 375 

unrepresented in any fossil record, known or yet undiscovered, be- 
cause successfully living fresh-water moUusk species seldom undergo 
fossilization. They usually enter the fossil record only when a 
habitat change brings about the extinction of populations or species. 
On the other hand, the present geographic distribution of the Thiaridae 
argues for their greater ability to spread across oceanic areas on island 
stepping stones without the necessity for any continuous "land 
bridges." 

There is one important taxonomic problem present in the Thiaridae 
that has not often been acknowledged in the study of these or any 
other parthenogenetic animal species. We know by observation the 
great variability of individuals within the species of the dioecious 
type such as the Pleuroceridae. This variability is the greatest 
single factor contributing to confusion in past and present studies of 
these fresh-water shells. The variability of individuals of the related 
Thiaridae may be either increased or decreased because of their 
reduction to unilateral ancestry. What we actually have in the 
entire family of the Thiaridae is the possibility of clone formation in 
every existing population of these animals. These is no biparental 
or cross-inheritance control or check on individual variation in their 
present parthenogenetic state. Wise indeed is the scientist who can 
tell whether a clone is a species or not, and be right every time, in the 
case of the Thiaridae. 

Family Thiaridae (in the Americas) 

The earliest generic name for any of the indigenous American 
species of this family is Aylacostoma Spix (1827, p. 15, pi. 8). This 
manuscript name of Spix was first published in the synonymy of both 
Melania tuberculata Wagner (= Aylacostoma tuberculatum Spix) and 
Melania scalaris Wagner ( = Aylacostoma glabrum Spix) in connection 
with the species description, and also on plate 8 (Spix, 1827) without 
the indication of synonymy. This is in the first edition (Munich) of 
plates, not the second edition (Leipsic). As a manuscript name of 
Spix, published in direct connection with two validly described and 
figured species, Aylacostoma is valid and available. The genotype, 
by subsequent designation by Morrison (1952, p. 8), is Melania 
scalaris Wagner (= Aylacostoma glabrum Spix). This species, as 
comparison of the figures will demonstrate, is that form called behnii 
by Reeve (1859-1861, pi. 2, fig. 8) in his 1860 monograph on Hemisinus. 

Swainson (1840, pp. 200, 341) was next to furnish a name for Ameri- 
can Thiaridae. Hemisinus is his generic name given to the species 
lineolata Gray from Jamaica. The later emendations, Aulacostoma 
Agassiz 1846 and Semisinus Crosse and Fischer 1885, are both 



376 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

unnecessary and invalid under present international rules of nomen- 
clature. Basistoma Lea (1852, p. 295) is exactly equivalent biologi- 
cally to Aylacostoma, being based on the same type of shell from 
South America. Verena H. and A. Adams (1854, p. 308) differs from 
the typical Aylacostoma group in shorter shell outline and in sculpture. 

Genu8 Cubaedomus Thiele, 1928 

This genus (Thiele, 1928, p. 401), based on the species hrevis Orbigny 
from Cuba, is the only American group clearly and generically distinct 
from Aylacostoma on shell characters. The shell is ovate-conic, with 
a trace of nodulous coronate sculpture on the upper slope or shoulder 
of the body whorl. There is no columellar notch or sinus in the 
aperture, which is evenly rounded below (anteriorly). The operculum 
is paucispiral. Animals of this species, dried in the shells, were 
recently boiled up in water (softened) and personally examined. 
These specimens (USNM 407991) were collected by Jaume from Eio 
"Los Cayos", Bahia Hondo, Pinar del Rio, Cuba. The mantle is 
fringed. The embryonic young are relatively large, and there were 
only two in the subhaemocoelic brood pouch in the neck region at one 
time in the material examined (pi. 11, fig. 19). Proof that this 
genus belongs to the family Thiaridae gives us another excellent 
example of concurrent and convergent evolution. Both PUsbry (1893) 
and Aguayo (1944, p. 69) have previously considered the species 
hrevis Orbigny of Cuba congeneric with {" Potamanax" = ) Pachychilus 
(Oxymelania) pilsbryi (Martens) of the Guatemala region, a member 
of the Pleuroceridae, because of the almost identical shell character- 
istics of the two species. 

Genus Aylacostoma Spix, 1827 

Subgenus Hemisinus Swaiason, 1840 

Genotype: {Melania lineolata Gray =) Aylacostoma (Hemisinus) 
lineolatum (Gray) 1828. 

This group of elongate-ovate or ovate-conic shells, almost completely 
lacking macroscopic spiral sculpture, includes one or two species from 
Cuba, the genotype from Jamaica, and a few other species of Central 
and South America from Panam^ to Ecuador and Peril. 

Subgenus Aylacostoma Spix, 1827, sensu stricto 

Genotype: (Aylacostoma glahrum Spix=) Aylacostoma (Aylacostoma) 
scalare (Wagner) 1827. 

This typical group includes one species from the Atlantic side of 
Central America, one species from the Pacific drainage in the Dari6n 
region of Panamd, and numerous species from the Magdalena, 
Orinoco, Amazon, and southern Brazil regions of South America. 



OLD AND NEW WORLD MELANL\NS — MORRISON 377 

These shells have spiral sculpture more or less prominent on most of 
of the whorls, and in some of the species there is a peculiar swelling 
or expansion of the body whorl of adults. The female reproductive 
anatomy and embryos (from the brood pouch) of the Central American 
species Aylacostoma ruginosum (Morelet) were figured by Crosse and 
Fischer (1892, pi. 49, fig. 10). In these figures the brood pouch 
relation is not clearly jndicated; it was not fully understood to be 
a separate adjunct to the primary female reproductive system. 
Hinkley (1920, p. 47) also has recorded the reproduction and ecology 
of this species. He says: 

Their trail was made by burrowing instead of crawling on the surface as with 
other forms. They burrowed somewhat like a mole, and often the little mole-like 
ridge could be followed quite a distance, and the moUusk found working under 
cover. 

It is a viviparous genus. When cleaning these shells the embryos run from one 
to three to the individual. None were noticed with more than three. 

Hinkley's observations on the embryos are corroborated by dried 
animals, received with the shells (USNM 218018) from Hinldey, 
which show two large young in situ in the brood pouch. These 
specimens are from Lake Ysabal, Jocolo, Guatemala. 

Subgenus Longiverena Pilsbry and Olsson, 1935 

Genotype: {Aylacostoma tuberculatum Spix, 1827 =Melania tubercu- 
lata Wagner, 1827, not Melania tuberculata (Miiller) 1776=) Ayla- 
costoma (Longiverena) tuberculatum Spix, 1827, by subsequent 
designation by Morrison, 1952. 

This subgeneric group of Aylacostoma species possesses prominent 
longitudinal sculpture in addition to the spiral lirae present on the 
shells of Aylocostoma, sensu stricto. In this way, the upper spire 
whorls may greatly resemble the corresponding portion of the shell 
of some species of Doryssa described from the same general region. 
More study is required for proof, but this general resemblance of 
Doryssa and Longiverena shells may be another outstanding example 
of concurrent and convergent evolution of the famiUes Pleuroceridae 
and Thiaridae. 

Pilsbry and Olsson (1935, p. 11) did not formally designate any of 
the originally included species, recent or fossil, as genotype, so we 
designated the living species tuberculatum in order that the animal 
characters of the group may be used to correctly place this unit in 
the total biological picture. 

Subgenus Verena H. and A. Adams, 1854 

ii£ Genotype: (Melania crenocarina Moricand=) Aylacostoma (Verena) 
crenocarina (Moricand). 



378 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

This fourth South American group, Verena, has been considered 
distinct by most authors. Its principal known distinguishing char- 
acter is the possession of strong, undulate or scalloped spiral ridges 
on the post-nuclear whorls. Dr. Olsson of the Academy of Natural 
Sciences of Philadelphia has recently found species in the fossil record 
from the upper Amazon region that prove the existence of Verena in 
that area for considerable time. These fossil species possess shells 
with sculpture identical in type to that of crenocarina (Moricand), 
but, unhke that living species, show shell outlines more or less com- 
pletely transitional to the high-spired outline of Aylacostoma, sensu 
stricto. 

Genus Tarehia H. and A. Adams, 1854 

One Asiatic species, probably Tarehia lateritia (Lea) from the 
Hawaiian Islands, has recently been introduced to American waters 
with aquarium plants. This species was first discovered living feral 
in Lithia Sulphm* Springs, near Tampa, Fla., in December 1947 by 
C. W. Cooke of the U. S. Geological Survey. It was not there in 
March 1940 on the occasion of an earlier complete collection of mol- 
lusk species by Dr. Cooke from that spring. This kind of introduction 
is highly dangerous to public health because this Asiatic type of snail 
is one of the primary intermediate host species of the human lung 
fluke (Paragonimiasis). 

Family Thiaridae (in Asia) 

Genus Thiara Roding, 1798 

This genus (Roding, 1798, p. 109) was not originally monotypic. 
It included two valid specific names, amarula (Linnaeus) 1758 (Chem- 
nitz, 1786, pi. 134, figs. 1218, 1219) and cancellata Rodmg 1798 
(Chemnitz, 1786, pi. 134, figs. 1220, 1221), as well as three nomina 
nuda. Hermannsen (1849, p. 576) did not explicitly state that 
amarula Linnaeus was the type. In Latin, he said it was a genus "of 
the type of." However, Brot (1874, p. 7) formally designated amarula 
the genotype. In the same monographic work he named two other 
groups which are synonyms of Thiara. Tiaropsis Brot (1874, p. 7), 
based on the genotype species winteri von dem Busch (1842, p. 1, 
pi. 1, figs. 1,2), and Plotiopsis Brot (1874, p. 7), based on the genotype 
species balonnensis Conrad (1850, p. 11), both possess the same shell 
characters as Thiara, differing principally in height of spire and size 
of shell. At present these differences are only considered of specific 
value. Another recently created synonym is Pseudoplotia Forcart 
1950, with the species scabra Miiller 1776 as originally designated 
genotype. 



OLD AND NEW WORLD MELANIANS — MORRISON 379 

Subgenus Thiara Roding, 1798, sensu stricto. 

Genotype: Thiara (Thiara) amarula (Linnaeus), 1758. 

The animals of Thiara spp. from Szechwan, China, Mindanao, 
Philippine Islands, and Okinawa, Ryukyu Islands, have been exam- 
ined by me and found to possess the reproductive structures described 
above as distinguishing characteristics of the family (pi. 11, figs. 9, 11). 

Subgenus Setaeara Morrison, 1952 

Genotype: (Thiara cancellata Roding 1798=) Thiara (Setaeara) 
cancellata Roding 1798. 

The genotype has been known for many years under the name of 
setosa Swainson because the far earlier name of Roding has not 
hitherto been critically examined. These shells possess stronger 
spiral sculpture than those of Thiara, sensu stricto, and the "crown" 
of spines on the shoulder of the whorls is continued upward as a set 
of cuticular setae, easily broken off, and almost always missing from 
imperfect shells. This group of only a few species was proposed as 
a subgenus (Morrison, 1952, p. 8) imtil animal characters are found 
that confirm its separation from Thiara, sensa stricto, by means of 
the shell. 

Genus Tarehia H. and A. Adams, 1854 

Genotype: (Melania semigranosa von dem Busch =) Tarehia semi- 
granosa (von dem Busch) 1842, by subsequent designation by Morri- 
son, 1952. 

Reexamination of the type designation of this genus by Brot (1874, 
p. 7) shows it to be completely inoperative. His statement that 
^'granijera" is the type refers to Melania granifera von dem Busch, 
the only such specific name originally included by H. and A. Adams. 
To the best of my knowledge, T. granifera H. and A. Adams is still a 
nomen nudum. This nomen nudum is proven not to be the same 
as Melania granifera of Lamarck by H. and A. Adams, who listed 
Lamarck's specific name separately and distinctly as a member of 
the genus Plotia. One of the valid, originally included species was 
designated as genotype (Morrison, 1952, p. 8) in order to correctly 
and legally fix Tarehia upon the group of thiarid snails to which it 
has been restricted and applied since the time of Brot (1874). 

The animals of T. lateritia (Lea) from China, of T. granifera 
(Lamarck) from the Philippines, and, more recently, of the Tarehia 
colony accidentally introduced to Lithia Sulphur Springs, near 
Tampa, Fla., have been examined for the reproductive characters 
and proven thiarid. 



380 PROCEEDINGS OF THE NATIONAL MUSEUM vol. io3 

Genus Sermyla H. and A. Adams, 1854 

Genotype: (Melania mitra Dunker =) Sermyla tornatella (Lea) 1850. 

Kecent examination of the animals of S. kowloonensis S. F. Chen 
1943 and of S. mauiensis (Lea), from China and the Hawaiian Islands 
respectively, proves this genus typically thiarid, as most authors have 
considered it. These shells possess arcuate, longitudinal sculpture on 
the upper (posterior) portion of the whorls, strongly contrasting with 
the spiral lirae below (anterior). They are relatively short-spired. 

Genus Stenomelania Fischer, 1885 

Genotype: (Melania aspirans Hinds =) Stenomelania aspirans 
(Hinds). 

Stenomelania Fischer (1885, p. 701) is a thiarid genus of elongate 
shells with the spire sometimes exceedingly attenuate and sharp- 
pointed, when not eroded away. Members of this genus possess the 
brood pouch typical of the family, that holds the eggs until they 
develop into veligers. Seshaiya (1940, p. 331) has shown that these 
snails, living in estuarine habitats, release the embryonic young into 
the water in the veliger stage. There is thus a free-swimming larval 
stage in the life history. These observations recently have been 
corroborated in the case of certain Philippine species of the genus by 
R. T. Abbott. The species M. aspirans is the genotype by monotypy. 
Another species, M. hastula (Lea), is the genotype by original desig- 
nation of the synonymous generic name Radina of Preston (1915, 
p. 10). 

j; ., Genus Melanoides Olivier, 1804 

Genotype: (Melanoides fasciolata Olivier =) Melanoides tuber culata 
(MiiUer) 1776. 

The shells of Melanoides are elongate-conic, of a number of regularly 
increasing whorls. The spire is usually twice the length of the 
aperture, or more. The sculpture is smooth to finely or coarsely 
tuberculate, in different species, without any material difference 
between the younger and older portions of the shell. The operculum 
is clearly paucispiral, with the nucleus so near the base that in some 
cases it may approach the subspiral condition. 

A personal check on available material in the U. S. National 
Museum collections has shown that the following species belong to 
Melanoides, agreeing exactly in the thiarid type of ovoviviparous, 
parthenogenetic reproduction: M. chinensis (Nevill), China; M. 
suijuensis S. F. Chen, Szechwan, China; M. turriculus (Lea), Leyte, 
Philippine Islands ; AfeZanoicZes sp., Rota, Marianas Islands; M. vainaja 
(Gould), Ofu, Samoa; and M. indeiinata (Lea), Oahu, Hawaiian 
Islands. 



OLD AND NEW WORLD MELANIANS — ^MORRISON 381 

Genus Sulcospira Troschel, 1857 

Genotype: (Melania sulcospira Mousson=) Sulcospira sulcospira 
(Mousson). 

Recorded as "also viviparous," this group must be studied further 
before its true relationships to the other genera from the Malayan 
region can be proved. Troschel (1857, p. 114) has recorded the 
radular characters; the shell appears similar to certain species of 
Brotia, to Tylomelania, or to immature individuals of Balanocochlis. 
Until the animal characters are reexamined, Sulcospira may be left 
tentatively in the "Melanoides complex." 

Genus Balanocochlis Fischer, 1885 

Genotype: (Melania glans von dem Busch=) Balanocochlis glans 
(von dem Busch) 1842. 

Philippine animals of this species, collected by R. T. Abbott at San 
Ramon, Mindanao (USNM 543951), have recently been examined. 
These animals prove the genus to be thiarid. There is a minute 
brood-pouch pore bordered with "flaps," similar to that figured for 
Melanoides turriculus (pi. 11, fig. 14), low on the right side and close 
to the fringed mantle edge. Apparently there are many hundreds of 
minute eggs in the brood pouch. Either they develop as numerous, 
small, equal-sized young shells as in certain Melanoides species, or 
they may be released in the veliger stage for a free-swimming larval 
period, as is known for the genus Stenomelania. The material ex- 
amined, which contained only the eggs, is not conclusive on this point. 
In any case, the shell and opercular characters are distinct enough 
from any other thiarids known to maintain the full generic separation 
of Balanocochlis. The earlier name Melania inermis Lesson 1830 was 
preoccupied by Gray in 1825; the later name M. siccata von dem 
Busch 1843 probably represents young of the species Balanocochlis 
glans. 

Genus Tylomelania P. and F. Sarasin, 1898 

Genotype: Tylomelania neritiformis P. and F. Sarasin. 

This subpaludomiform group of shells only appears as a separate 
development of shell outline and proportions ; they have the same type 
of "paleomelanian" operculum and the same general radular char- 
acters as do Antimelania and Brotia. The animals of all the groups 
must be completely and carefully reexamined before the genera (and 
the distinctions of the genera) of the "Melanoides complex" of the 
Thiaridae can be properly organized or understood. Shell, radular, 
and opercular characters of this group were carefully recorded by the 
Sarasins (1898, p. 52, pi. 4, figs. 56, 57; pi. 8, figs. 110, 111; pi. 9, fig. 
115), but they did not examine the details of the reproduction of 
Tylomelania. 



382 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loa 

Genus Antimelania Crosse and Fischer, 1892 

Genotype: (Melania variabilis Benson 1836 (not Melania variabilis 
Defrance 1823)=) Antimelania costula (Rafinesque) 1833. 

As the well-known but preoccupied and preceded name for the 
genotype implies, the shell sculpture of members of this genus is 
variable. These shells are similar to Melanoides but usually larger in 
size, "with the sculpture bolder when present. The operculum is 
paleomelanian, often with about 6 turns in contrast to that of 
Melanoides with about 2 turns. The apex or nuclear whorl of these 
shells is perfectly symmetrical in uneroded individuals. The aperture 
is more-or-less regularly rounded at the base, without a forward 
projecting rounded angle as in Brotia. 

Specimens of Antimelania sorinianxL (Heude) collected in 1940 by 
Dr. H. T. Chen at Tai-Mo-Shan, Kwangtung, China, possess a paleo- 
melanian operculum (with large nucleus) as in Heude's figures (1890, 
pi. 41, figs. 6, 6a). They were all ovo viviparous, parthenogenetic 
females containing hundreds of minute young in the brood pouch, 
which is enormously expanded into the body cavity behind the head. 
They possessed a narrow genital groove terminating abruptly on the 
right side of the foot, without any trace of the pleurocerine type of 
egg-laying sinus. 

Apparently Pilsbry and Bequaert (1927, p. 300) did not know that 
the specific name variabilis was preoccupied when they designated it 
the genotype of Antimelania (Crosse and Fischer, 1892, p. 313). A 
search for the correct name to use in replacement led to the discovery 
that costula Rafinesque (1833, p. 166) is clearly recognizable as identi- 
cal to Benson's variabilis. 

Moore (1899a, p. 161) has recorded the brood-pouch ovoviviparity 
of ^^Melania" episcopalis (Lea). His figure (pi. 14, fig. 13) of the 
female Antimelania episcopalis (Lea) agrees with our observations on 
the genus Antimelania. The figure given by him of a "male" animal 
of this same species is either a nongravid or nonbreeding female of the 
same species, or a female of the pleurocerine genus Oxytrema with only 
the genital groove and egg-laying pit in the side of the foot. Moore 
(1899a, p. 163) mentions the presence of males in Tanganyicia and 
'^Melania," but nowhere proves it. The presence of any males in 
this family is still not proved. 

Genus Brotia H. Adams, 1866 

Genotype: (Melania pagodula Gould 1847 (not Melania pagodulus 
Reeve 1860)=) Brotia pagodula (Gould) 1847. 

Shells of this genus are elongate-conic, of variable sculpture, and 
with the columellar angle of the basal lip of the aperture produced 



OLD AND NEW WORLD MELANIANS — MORRISON 383 

slightly forward. The operculum is of the paleomelanian type, of 
many (6 or 8) turns, and, in the species seen, is considerably smaller 
than the aperture. The perfect apex of the shell (nuclear whorl) is 
asymmetrical. Specimens of the animals of Brotia baccata (Gould) 
and the adult and embryonic shells of three other species of Brotia 
from the Salween River of Thailand and Burma have been personally 
examined. In the species baccata there is a well-developed, deep 
genital groove do\vn the right side of the foot that abruptly turns in- 
ward to become the entrance (or "birth pore") of the brood pouch. 
This brood pouch is very large, crowded with a great many relatively 
small young of uniform size. The single character of the embryonic 
shell, or the apex (when perfect) of the adult shell, that characterizes 
Brotia is the asymmetry. It appears as if each shell in its development 
is interfered with by a yolk sac (?) in the position of the apex to keep 
that apex soft, while the rest of the embryonic whorls develop nor- 
mally and symmetrically, and then the apex collapses to a line below 
the curve of spiral symmetry. This may very well be connected with 
a secondary development in the brood pouch to insure a "nurse" type 
of nutrition of the young. 

Synonyms of Brotia include Acrostoma Brot 1874, Brotella Rovereto 
1899, Paracrostoma Cossman 1900, with the genotype hugeli Philippi, 
and Wanga S. F. Chen (1943, p. 21), based on the species Melania 
henriettae Gray (1834, pi. 13, fig. 2). In this connection it should be 
mentioned that the original figure of henriettae is poorly drawn and 
does not show the generic character of the slightly produced columellar 
angle of the aperture. This fault is corrected in the figure of the 
holotype published by Yen (1942, p. 204, pi. 15, fig. 66). In fact it is 
possible that the originally published locality of henriettae ("China") 
is at fault, and this may be the same species as baccata (Gould) of the 
Salween River and some of its tributaries. 

Genus Fijidoma Morrison, 1952 

Genotype: (Fijidoma laddi Morrison 19 52)= Fijidoma maculata 
(Mousson) 1865, by original designation. 

Fijidoma Morrison (January 1952) antedates by only a few months 
the synonymous generic name Veloplacenta Hubendick (August 1952), 
which was based on the same species of thiarid snail. These fresh- 
water snails from Viti Levu in the Fiji Islands, superficially very 
similar in shell outline to those of the southeast Asiatic genus Paludo- 
mus, were named in allusion to that similarity. They have rather 
smooth, subglobose, neritiform shells with a few rather rapidly in- 
creasing, well-rounded whorls separated by a deep suture. The shell 
has fine spiral sculpture and regular flammules or rows of squarish 
dots (interrupted flammules) of red in the ground color. The aper- 



384 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 103 

Explanation of Plate 11 

1, Melanopsis dufourii (Ferussac), female, from the right side showing egg-laying groove 

and the "ovipositor" in pit on right side of foot, from Elche, Alicante Province, Spain 
(USNM 515845). 

2, Oxytrema canaliculatum undulatum (Say), female, showing egg-laying groove and pit in 

right side of foo